Pak. J. Bot., 43(6): 2769-2785, 2011.
FURTHER INSIGHTS AND NEW COMBINATIONS IN AYLOSTERA (CACTACEAE)
BASED ON MOLECULAR AND MORPHOLOGICAL DATA
STEFANO MOSTI1,3, NADEESHA LEWKE BANDARA2 AND ALESSIO PAPINI1,3*
1
Dipartimento di Biologia Evoluzionistica; Università degli Studi di Firenze Via La Pira, 4-50121 Firenze
2
The John Bingham Laboratory, NIAB. Huntingdon Road, Cambridge CB3 0LE, UK.
3
Centro Studi Erbario Tropicale (CSET) of the University of Florence, Via La Pira, 4-50121 Firenze
Pubblicazione del Centro Studi Erbario Tropicale n 119.
Abstract
The Genus Rebutia K. Schum. is a taxonomically complex genus of Cactaceae subfamily Cactoideae. This genus was
subjected to splitting and lumpering treatments through the years. Molecular data revealed that Rebutia sensu lato must be
divided in a clade corresponding to former Rebutia section Rebutia (Rebutia sensu stricto), related to the clade formed by
Weingartia/Cintia/Sulcorebutia. Only this clade corresponds to genus Rebutia in our results. The other sections of Rebutia s.
l. (Aylostera, Digitorebutia, Cylindrorebutia) cluster together to form another clade not directly related to Rebutia s. str. For
priority reason this clade is recombined as genus Aylostera Speg. An analytical key is provided, to identify genera Rebutia
K. Schum., Aylostera Speg., and Weingartia Werderm. (including Sulcorebutia Backeb. and Cintia Knize & Riha) and in
Aylostera, at infrageneric level, the subgenera Aylostera and Mediolobivia. Further investigations are needed to assume
taxonomic decisions about the clade Weingartia/Sulcorebutia/Cintia that, as a whole, should be assigned to genus
Weingartia Werderm. A list of taxa belonging to genera Rebutia K. Schum. and Aylostera Speg., is provided. In this
treatment the necessary combinations following the separation of Aylostera as a genus autonomous from Rebutia are
proposed. The ATPB-rbcL IGS fragment revealed to be enough variable to be used for Barcoding of species among
Cactaceae. All species here considered are in CITES appendix II, with frequent determination difficulties.
Introduction
The genus Rebutia K. Schum., was described in 1895.
Since the beginning the genus was subjected to various
rearrangements. This tendency lead to decisions
sometimes extreme, both on a “splitter” and on a
“lumper” point of view.
An example of a “lumper” treatment was the
rearrangements that lead, quite recently, to the fusion of
Rebutia with the genera Sulcorebutia Backeb. and
Weingartia Werderm (Hunt & Taylor, 1990; Hunt, 1999;
2006). On the contrary an example of a “splitter” point of
view was in the previous treatments as that by Backeberg
(1966), who subdivided Rebutia in three genera: Rebutia
K. Schum. sensu stricto, Aylostera Speg., and
Mediolobivia Backeb., with the allied genera Sulcorebutia
and Weingartia.
An intermediate treatment was carried out by Donald
(Donald, 1975a; Donald & Brederoo, 1975), who finally
decided to divide the genus Rebutia K. Schum. (excluding
Sulcorebutia and Weingartia) into five sections without
subgenera: Rebutia K. Schum. (including subsections
Rebutia Buining & Donald and Mediorebutia Buining &
Donald), Aylostera Speg., Digitorebutia (Frič & Kreunz.
ex Buining) Buining & Donald, Cylindrorebutia Buining
& Donald and Setirebutia Buining & Donald. This point
of view was accepted also by Mosti (1999).
Moreover, Donald (1975b) and Donald & Brederoo
(1976a; 1976b; 1976c; 1976d; 1977a; 1977b; 1978a;
1978b) revised four of the five sections. The fifth section,
Digitorebutia, was revised by Mosti (1999; 2000a;
2000b).
Rausch, the most important discoverer of Rebutia
sensu lato, since 1986 treated Digitorebutia and the few
species belonging to Cylindrorebutia and Setirebutia
under the genus Lobivia Britton & Rose (Rausch, 1986).
Moreover, this last genus was recently included in
Echinopsis Zucc., by Hunt & Taylor (1990). Rausch’s
opinion about the inclusion of part of Rebutia in Lobivia
and, transitively, in Echinopsis, is not currently accepted
(Sida, 1997; Hunt, 1999; Mosti, 1999; Anderson, 2001).
A summary about the history of the main taxonomic
treatments about Rebutia and allied genera is given in
Table 1.
Table 1. History of the taxonomic treatment of genera: Rebutia, Aylostera, Weingartia, Sulcorebutia and Cintia.
Backeberg (1966)
Donald (1975)
Rowley (2009)
Mosti & Papini (2011)
Genus Rebutia K. Schum.
Genus Rebutia K. Schum Genus Rebutia K. Schum Genus Rebutia K. Schum.
sect. Rebutia
subgen. Rebutia
Genus Aylostera Speg.
Genus Rebutia sect. Aylostera Genus Aylostera Speg. subgen. Genus Aylostera Speg. subgen.
Aylostera
Aylostera
Genus Mediolobivia Backeb. Genus
Rebutia
sections Genus Aylostera Speg. subgen. Genus Aylostera Speg. subgen.
Digitorebutia,
Mediolobivia
(Backeb.) Mediolobivia (Backeb.) Rowley
Cylindrorebutia
and Rowley
Setirebutia
Genus Weingartia Werderm. Genus Weingartia Werderm.
Genus
Rebutia
subg. Genus Weingartia Werderm.
Weingartia
(Werderm.) including genera Sulcorebutia
Backeb. and Cintia Knize &
Rowley
Riha
Genus Sulcorebutia Backeb. Genus Sulcorebutia Backeb.
Genus
Rebutia
subg.
Sulcorebutia (Backrb.) Brandt
*
Corresponding author E-mail: alpapini@unifi.it
2770
Digitorebutia, Cylindrorebutia and Setirebutia, or as
a whole the subgenus “Mediolobivia” sensu Pilbeam
(1997), as the other rebutias, should be considered
separated from the Lobivia/Echinopsis group because of
essential morphological differences, including indehiscent
fruits and stamens disposed in only one series which are
present only in Rebutia s. l.
Ritz et al., (2007) proposed, on the basis of molecular
data, to separate Rebutia group I, that is, the few analysed
by them corresponding to genera Aylostera Speg. (R.
deminuta, R. fiebrigii and R. pseudodeminuta) and
Mediolobivia Backeb. (R. einsteinii, R. pygmaea and R.
steinmannii) after Backeberg (1966), from Rebutia group
II (R. minuscula, and R. padcayensis) corresponding to
genus Rebutia K. Schum s. str. in Backeberg’s (1966)
system.
The species belonging to group II resulted closely
related to Sulcorebutia/Weingartia (clade E) in Ritz et al.,
(2007). The proposal to consider Rebutia s. str. separated
from Aylostera and Mediolobivia was based both on
biomolecular data and enforced by morphological
characters:
Rebutia
sensu
Backeberg
(1966)
(corresponding to Rebutia s. str.) own in fact hairless
pericarpels, while on the contrary, Aylostera and
Mediolobivia have hairy and bristly pericarpels. This
proposal, partially corresponding with Backeberg’s (1966)
idea to consider Rebutia s. l. (excluding Sulcorebutia and
Weingartia) as splitted in more than one genera, was not
followed by a formal taxonomical recombination by Ritz
et al., (2007).
On the basis of Ritz et al., (2007) findings, Rowley
(2009) proposed a classification of Rebutia s.l. separating
Rebutia s.str., including Weingartia and Sulcorebutia as
subgenera and the genus Aylostera was divided in the two
subgenera Aylostera and Mediolobivia. Aylostera Speg.,
has in fact priority with respect to Mediolobivia Backeb.,
to Digitorebutia Frič & Kreuz. ex Buining and to
Setirebutia Frič and Cylindrorebutia Frič & Kreuz.. These
last three names were never described as genera with a
Latin diagnosis.
The aim of our investigation was to verify and
enforce the data obtained by Ritz et al., (2007), that
included a too limited number of species of Rebutia, by
adding new molecular data on taxa belonging to genus
Rebutia s.l., that is sensu Buining & Donald (1963; 1965)
(hence excluding gen. Sulcorebutia and Weingartia), and
correlating them with morphological characters, in order
to clear the complex taxonomical classification of this
group.
A secondary aim was to test a single molecular
marker, the ATPb-rbcL spacer as potential Bar-coding
marker for Cactaceae. All species here considered are in
CITES appendix II, with frequent determination
difficulties: the possibility of using a molecular marker to
identify them with certainty might be of interest for
conservation purpose.
Materials and Methods
Plant material: The plants used in the analysis are
cultivated by the Botanical Garden of Florence “Giardino
dei Semplici” and collection S. Mosti and were obtained
via vegetative propagation from the samples collected by
older researchers in the wild before the insertion of
Cactaceae in Appendix 1 and 2 of CITES rules. The
original samples were those used for species description.
The indicated field numbers are those used by the
STEFANO MOSTI ET AL.,
collectors in the wild. Our samples are deposited in the
Tropical Herbarium of Florence (FT). The taxa used for
the molecular analysis are known often for a single or few
populations in Aylostera, Mediolobivia and Rebutia. The
same can be said in other genera such as Echinopsis.
We added the following taxa to Ritz et al., (2007)
data set (DNA Genbank accession at the end of each
sample): Rebutia wessneriana Bewer. subsp. berilloydes
(Buining & Donald) Donald, f. n. (field number) Rausch
819, cult S. Mosti 2011 (FT!), GU084402; R. kariusiana
Wessner, s.n., cult. S. Mosti 2011 (FT!), GU084394; R.
fabrisii Rausch var. aureiflora Rausch, f. n. Rausch 687,
cult. S. Mosti 2011 (FT!) belonging to sect. Rebutia even
if Pilbeam (1997) links R. fabrisii to Aylostera (for this
species we chose this yellow flowered variety with a
single population, but even a red flowered population is
known.); GU084401; Rebutia donaldiana A.B. Lau &
G.D. Rowley, f. n. Lau 348, cult. S. Mosti 2011 (FT!),
GU084399; R. muscula F. Ritter & Thiele, f. n. Ritter 753,
cult. S. Mosti 2011 (FT!) belonging to subgen. Aylostera,
GU084400; Rebutia haagei Frič & Schelle, f. n. Rausch
35, cult. S. Mosti 2011 (FT!) (R. haagei is a species
characterized by a slight variability in some populations.
The variability regards the shape of the petals with a more
or less evident mucrone and some slight variation in the
color of the flowers), GU084396; R. christinae Rausch, f.
n. Rausch 492a, cult. S. Mosti 2011 (FT!), GU084398; R.
friedrichiana Rausch, f. n. Rausch 646, cult. S. Mosti
2011 (FT!) belonging to subgen. Mediolobivia (Backeb.)
Rowley, GU084397.
Moreover we included in the analysis some samples
of taxa that, they do not even belong to genus Rebutia s.l.,
but are noteworthy for an analysis about south american
mountain cacti. All these taxa belong to genus Echinopsis
s. l.: Lobivia haematantha (Speg.) Britton & Rose var.
haematantha (= Echinopsis haematantha (Speg.) D.R.
Hunt), s.n., cult. S. Mosti 2011 (FT!), GU084392.;
Lobivia (Chamaecereus) silvestrii (Speg.) G.D. Rowley
(= Echinopsis chamaecereus H. Friedrich & Glaetzle),
s.n., cult. A. Papini 2011 (FT!), GU084395;
Acanthocalycium glaucum F. Ritter (= Echinopsis
glaucina H. Friedrich & G.D. Rowley), s.n., cult. S. Mosti
2011 (FT!), GU084391 and Echinopsis melanopotamica
Speg. (recently included in Echinopsis leucantha (Gillies
ex Sal-Dyck) Walp.), s.n., cult. S. Mosti 2011 (FT!),
GU084393.
E. melanopotamica and L. haematantha are,
respectively, an Echinopsis s.str. and a Lobivia s.str. L.
silvestrii is a species of interest because of its
morphology, resembling to that of Rebutia for its thin and
lengthened stem, for its short and fragile spines and the
cespitous habit. Acanthocalycium glaucum, finally, is a
representative of the only genus of Echinopsis s.l. not
included in Ritz et al., (2007) analysis.
All new samples introduced in the analysis are
endemic of North Argentina/South Bolivia living at high
altitude (more than 3000 m above sea level).
For the phylogenetic analysis we used the same
outgroups used by most recent molecular studies
(Nyffeler, 2002; Ritz et al., 2007. The ingroup taxa
represent South American cacti of the ‘‘BCT clade’’ as
defined by Nyffeler (2002) with an emphasis on the
genera Gymnocalycium, Rebutia, Sulcorebutia, and
Weingartia.
Molecular data: We amplified the DNA fragment
ATPB-rbcL spacer, with the same primers used by Ritz et
NEW COMBINATIONS IN AYLOSTERA
al., (2007). This fragment showed the highest variability
among the three used by Ritz et al., (2007), that is more
than half of the informative substitutions and half of the
informative indels within the ingroup.
Genomic DNAs were isolated from flowers in buds
(3-4 days before anthesis) of individual plants using a
modified CTAB extraction protocol (Doyle & Doyle,
1990; tissue ground in sea-sand, 70% [v/v], isopropanol
substituted for the RNAase step). DNA concentrations
were estimated by gel electrophoresis on 1% agarose. We
used one sample for each accession.
ATPB-rbcL fragment sequencing and phylogenetic
analyses: Primer sequences for the 50 region of the atpBrbcL IGS were taken from Savolainen et al., (1994): ‘‘2’’
5'-GAAGTAGTAGGATTGATTCTC-3' and ‘‘10’’ 5'CATCATTATTGTATACTCTTTC-3', as suggested by
Ritz et al., (2007). The amplification was performed for
180 s at 958C; followed by 28 cycles of 30 s at 958C, 60 s
at 428C, and 120 s at 728C; and a final extension for 180
s at 728C. Single-banded fragments were purified and
directly sequenced in both directions by using the
amplification primers. Cycle Sequencing and the BigDye
Terminator Ready Reaction Kit (Applied Biosystems)
were used. Data were collected on ABI Prism 373A
automated gel reader in the laboratory CIBIABI of the
University of Florence. Resulting sequences were further
checked by eye with the software CHROMAS 2.3
(www.technelysium.com.au). A BLAST (Altschul et al.,
1997) search was performed to exclude the sequencing of
any contaminant organism. The boundaries of the
fragments were determined by comparison with
previously published sequences. We added our new data
on ATP-rbcL IGS to the previous alignment by Ritz et al.,
(2007), including there data on trnL-trnF and trnK-rps16
IGS sequences. The matrixes were combined with the
Python (Python version 2.6.4; Biopython 1.57) program
combinex1_0.py, written by one of the authors, A. Papini,
released under GPL licence and available at
www.unifi.it/caryologia/PapiniPrograms.html.
Optimal multiple alignment was obtained with
CLUSTALW 1.81 (Thompson et al., 1994) and checked
by eye. Parsimony analysis was performed with PAUP*
4.0b1 (Swofford, 1998). We used a successive weighting
approach (Farris, 1969) because of the high number of
trees obtained by a simple hsearch command. We used the
first 20000 trees to reweight characters (command
REWEIGHT) for a second hsearch analysis and so on
until, since the third run, the characters weight did not
change anymore.
At this point a heuristic search analysis was run with
TBR branch-swapping, multrees option on, addseq =
simple, ten randomised replicates., a hsearch analysis was
performed with a maximum number of saved trees of
65197 to reduce computational effort (50000 were
retained by Ritz et al., 2007). All characters were
weighted equally, and character state transitions were
treated as unordered. Gaps were treated as "simple indel
coding" after Simmons & Ochoterena (2000), coding
them with the software Gapcoder (Young & Healy, 2003).
This process codes indels as separate characters in a data
matrix, which is then considered along with the DNA base
characters in phylogenetic analysis.
A maximum likelihood (Felsenstein, 1981) search
was done as follows: we used Modeltest 3.06 (Posada &
2771
Crandall, 1998) to evaluate the likelihood of 56 different
models of sequence evolution on the basis of our data.
The likelihood ratio test option was used to compare
likelihood scores in a nested design. We used the most
likely model of evolution from Modeltest 3.06 as settings
in a maximum likelihood (ML) phylogenetic analysis in
PAUP. We used the obtained model to calculate the
likelihood value of the maximum parsimony trees.
Because of computational time problem with PAUP,
we performed a maximum likelihood analysis with the
GARLI package (Zwickl, 2006) that uses a stochastic
genetic algorithm-like approach to simultaneously find the
topology, branch lengths and substitution model
parameters that maximize the log-likelihood (lnL). The
package was used on a server provided by the Cipres
portal (Miller et al., 2009 for the site address).The indelsderived characters were excluded for the maximum
likelihood analysis.
Bootstrap (Felsenstein, 1985) resampling was
performed using setting search = faststep (with no TBR
branch-swapping because of computational time limits)
with ten random taxon entries per replicate and multrees
option in effect (with 10000 replicates) under parsimony
criterion.
MrMODELTEST 2.0 (Nylander, 2004) was used to
test the best model of sequence evolution (based on the
Akaike Information criterion, Akaike, 1974) to be used
with the program for Bayesian Inference MrBayes 3.4b4
(Huelsenbeck, 2001). For the indels-coded characters we
used the same evolutionary model used for restriction
sites that are coded as binary character states (as
implemented in MrBayes).
The Bayesian phylogenetic analysis was used for
assessing the robustness of tree topology and the support
for clades. The posterior probability of the phylogenetic
model was estimated using Markov chain Monte Carlo
(MCMC) sampling with the Metropolis-Hastings-Green
algorithm. Four chains were run, three heated and one
cold, for 106 generations and sampled every 100
generations. Following the analysis, the posterior
probabilities were checked in the output of Mrbayes to
estimate the number of trees that should be discarded as
"burn-in”. Stationarity was reached at approximately
generation 30,000, so the first 300 trees or “burn-in”
period of the chain were discarded. Phylogenetic
inferences are therefore based on those trees sampled after
generation 30,000. After the "burn-in" trees were removed
from the data set, the remaining trees were used to
produce a 50% majority-rule consensus tree (with PAUP)
in which the percentage support indicated a measure of
the Bayesian posterior probabilities.
To test the significance of the difference of less
parsimonious trees with respect to the most parsimonious
solution, the Templeton (Wilcoxon signed-ranks) test
(Templeton, 1983) was used as implemented in PAUP.
This test was used to evaluate the significance of an
alternative position of taxa of Rebutia s. l.
A matrix of the genetic distances among the different
species of Rebutia was calculated with PAUP and
presented in Table 2. The distances were obtained by
calculating the total difference (number of total
substitutions in the matrix of DNA sequences).
The trees were edited for better readability with
Dendroscope (Huson et al., 2007).
2772
STEFANO MOSTI ET AL.,
Table 2. Matrix of distances obtained by total difference (number of total substitutions in the matrix of DNA
sequences). First row: abbreviations of species reported in the first column.
sulc pse fie de pyg ste ein pad min fab kar wes haa fri chr don mus
Sulcorebutia steinbachii
0
4
30 31 39 20 34 39 24 15 18 18 26 26 27 30 28
Rebutia pseudodeminuta
0
0
0
12 11 12 10 12 0
0
0
0
0
0
0
0
Rebutia fiebrigii
0
13 32 31 28 26 33 17 19 19 21 21 22 16 15
Rebutia deminuta
0
25 19 31 30 31 16 17 17 12 12 13 6
4
Rebutia pygmaea CA08
0
4
28 37 38 27 28 28 14 14 15 27 26
Rebutia steinmannii
0
16 16 21 9
11 11 1
1
1
10 9
Rebutia einsteinii
0
30 32 16 18 18 19 19 20 19 19
Rebutia padcayensis
0
32 23 24 24 34 34 35 36 35
Rebutia minuscule
0
7
4
4
19 19 20 24 24
Rebutia fabrisii
0
4
4
15 15 14 17 17
Rebutia kariusiana
0
0
17 17 18 21 21
Rebutia wessnerana ssp.
0
17 17 18 21 21
beryllioides
Rebutia haagei
0
0
1
14 14
Rebutia friedrichiana
0
1
14 14
Rebutia christinae
0
13 13
Rebutia donaldiana
0
3
Rebutia muscula
0
Results
The total alignment consisted of 2866 characters, of
which 2335 resulting from nucleotides sequence
alignment of ATPb-rbcL, trnL-trnF and trnK-rps16 IGS
sequences, while other 531 characters were added as a
result of indels coding. Echinopsis atacamensis subsp.
pasacana and Lobivia haematantha showed a big
insertion in 384-492 position of the ATPb-rbcL
alignment.
The successive weighting and maximum parsimony
analysis yielded 65197 trees (restriction of retained trees
with MAXTREE=65197) with L=1089.233 (some
weights are between 0 and than 1); CI=0.872; RI=0.880.
The strict consensus tree (Fig. 1, Bootstrap support above
branches) had L = 1094.249; CI = 0.868; RI = 0.875.
The maximum likelihood analysis obtained with GARLIC
is shown in Fig. 2 (Bayesian support above branches).
GARLIC identified the following nucleotide substitution
rates: AC = 0.779, AG = 0.637, AT = 0.269, CG = 0.782,
CT = 0.327, GT = 1.000. This rates were quite different
from those obtained with MODELTEST particularly for
the A-C and A-T substitution rates.
Our analysis largely agreed with Ritz et al., (2007)
and for this reason same clade labels were used.
The clade E (Weingartia, Sulcorebutia, Cintia,
Rebutia II, Browningia) was confirmed in our analysis
both with the strict consensus tree (parsimony) and the
maximum likelihood tree but with Bayesian and Bootstrap
support lower than 50%. We added to Rebutia II R.
kariusiana, R. wessneriana and R. fabrisii. All these
Rebutia clustered together with 99% Bayesian Support
(BS), Decay Index (DI) = 4 and 90% Bootstrap support
(BO). This clade corresponds to Rebutia sect. Rebutia
sensu Donald (1975a). Within Clade E the two species of
Browningia resulted as Outgroup to the rest of the clade.
The two Browningia did not cluster together.
Keeping the two representatives of Browningia
together resulted in trees L=1113.60343 (1094.24857 the
strict consensus of the maximum parsimony trees) with
CI=0.853 and RI=0.858. The difference resulted
statistically significant after the Templeton test
(p=0.0068). Keeping together the whole clade E but
Browningia collapsing the node dividing Rebutia sect.
Rebutia from Weingartia, Cintia and Sulcorebutia,
resulted in trees L=1096.75376, CI = 0.866 and RI=0.873
and the difference resulted statistically significant after the
Wilcoxon test (but with p=0.0273, while statistical
significance is considered to be for p<0.05 in PAUP.
The clade D, corresponding to Rebutia I (Aylostera,
Mediolobivia) by Ritz et al., (2007), was monophyletic
with 94% Bayesian support BS and DI = 1 (Bootstrap
support lower than 50%). Representatives of subgen.
Mediolobivia (labelled with A on Figs. 1 and 2) formed a
clade with 94% BS and DI=3 excluding R. einsteinii that
clustered as outgroup to the rest of Mediolobivia in the
maximum parsimony analysis (Fig. 1) and as outgroup to
both Mediolobivia and Aylostera in the maximum
likelihood analysis (Fig. 2).
Moving Aylostera clade to the Rebutia s. str. clade
(that is keeping together the whole genus Rebutia s.l.,
both those from clade E and those from clade D) resulted
in trees L=1097.95757, CI=0.865, RI=0.872 and the
difference resulted statistically significant after the
Wilcoxon test (p=0.0225). Moving Rebutia s. str. to the
Aylostera clade resulted in trees of L=1096.79986,
CI=0.866, RI=0.873 with significant difference after the
Templeton test (p=0.0146). Noteworthy P values resulting
by keeping Rebutia s. str. together with Aylostera were
not far from the 0.05 of significance value.
NEW COMBINATIONS IN AYLOSTERA
2773
Fig. 1. Strict consensus tree of 65197 maximum parsimony trees (both ATPb-rbcL IGS sequence and derived indels data. Bootstrap
support (1000 replicates no TBR branch swapping on) above branches. Clade E = Weingartia, Sulcorebutia, Cintia, Rebutia II,
Browningia as in Ritz et al., (2007). A = Aylostera subgen. Aylostera; C = Aylostera subgen. Mediolobivia. The whole genus Aylostera
corresponds to letter D and Rebutia I after Ritz et al., (2007).
STEFANO MOSTI ET AL.,
2774
Fig. 2. Maximum likelihood tree obtained with GARLIC package. Bayesian support (indels treated with restriction site model) and
Decay indexes above branches. Labels as in Fig. 1.
The matrix of the genetic distances (total number of
substitution) among the different species of Rebutia
calculated with PAUP (Table 2) showed that the
sequences used in this analysis were useful to verify the
relationships among different species.
Discussion
In clade E Rebutia K. Schum. sect. Rebutia (Buining
& Donald, 1963, 1965; Donald & Brederoo, 1975)
clustered together with one representative of Browningia,
while the other Browningia resulted either as outgroup to
NEW COMBINATIONS IN AYLOSTERA
the rest of lade E or in uncertain position. In Ritz et al.,
(2007) Browningia candelaris resulted outgroup to
Rebutia s.str., with B. hertlingiana in uncertain position.
The position of Browningia prevented us to consider
Rebutia sect. Rebutia and the clade composed by
Weingartia+Sulcorebutia+Cintia as belonging to the same
genus as proposed by Rowley (2009), who did not
consider the position of Browningia. The consequence is
that the species belonging to Rebutia sect. Rebutia remain
the only representatives of genus Rebutia K. Schum.
This genus corresponds to Rebutia s.str., which is
those species inserted in Rebutia K. Schum. by Backeberg
(1966) or to sect. Rebutia of genus Rebutia K. Schum. in
the sense of Buining & Donald (1963, 1965), but without
Sulcorebutia/Weingartia/Cintia. These three genera are
probably to be considered as congeneric, Weingartia (for
priority reasons) as was indicated also by Hentzschel &
Augustin (2008), but to be considered separated by
Rebutia (Rebutia group II after Ritz et al., 2007), also
because of fundamental morphological characters
(Augustin et al., 2000 and our further studies): the areoles
are in eccentrical position and sunken on the higher
(adaxial) part of the tubercle in Weingartia/Sulcorebutia,
while they are more superficial and on the central (distal)
part of the tubercle in Rebutia. The scales of the flower
tube are broad and imbricated in Weingartia/Sulcorebutia;
they are more narrow and not imbricated in Rebutia.
Finally the spines are quite fragile and subtle in Rebutia,
and more robust and thick in Weingartia/Sulcorebutia.
Clade D resulted separated from clade E Rebutias.
This result confirmed Ritz et al., (2007) results also with
our larger species sampling. The Wilcoxon test confirmed
that our data prevent keeping all Rebutia together. The
consequence is that clade D should be considered a
separate genus by using the priority name Aylostera Speg.
This genus corresponds to subgen. Aylostera and the
subgen. Mediolobivia (Backeb.) Rowley (corresponding
to the sections Digitorebutia (Frič & Kreuz. ex Buining)
Buining & Donald, Cylindrorebutia Buining & Donald
and Setirebutia Buining & Donald of Rebutia K. Schum).
R. einsteinii resulted included in subgen. Mediolobivia in
the maximum parsimony analysis, while it was outgroup
to all the other accessions of the new genus Aylostera in
the maximum likelihood analysis. R. einsteinii is the only
representative of sect. Cylindrorebutia inserted in the
analysis, and its position in the maximum likelihood
analysis would be compatible with a treatment of the
genus as subdivided in sections. The sect. Setirebutia (not
inserted in the analysis since it is synonymized in our
treatment with Mediolobivia) is considered belonging to
genus Aylostera because of its hairy pericarpels.
Only a few chromosome counts are available for
Rebutia s.l. in our knowledge. Anyway Ross (1981)
observed that R. minuscula and R. violaciflora, both
belonging to Rebutia s. str., have n=11 chromosomes as
Sulcorebutia steinbachii (Rebutia steinbachii in Ross,
1981). These three species all belong to clade E. On the
contrary R. kupperiana and R. spegazziniana, both to be
assigned to clade D (Aylostera) had n=22. Hence these
few chromosome counts would confirm our molecular
results.
2775
Taxonomic treatment
The taxonomical consequences of our analysis lead to
a modified classification with respect to the recent
reappraisal by Rowley (2009) as reported underneath. The
new classification is preceded by an analytical key that
permits to identify genera Rebutia K. Schum., Aylostera
Speg., and Weingartia Werderm. (including Sulcorebutia
Backeb. and Cintia Knize & Riha) and in Aylostera, at
infrageneric level, the subgenera Aylostera and
Mediolobivia.
About the clade Weingartia/Sulcorebutia/Cintia we
did not take either taxonomic decisions or new
recombinations since the knowledge about this group
require further investigation, also about taxa delimitation.
The monotypic genus Cintia Knize & Riha, despite its
phylogenetic position closely related to Weingartia and
Sulcorebutia, as observed in the molecular analysis (Ritz
et al., 2007 and our data), shows peculiar morphological
characters (autoapomorphies) such as lack of spines and
very short (or even absent) floral tube. The flower appears
sunken in the areole.
We report also a description of genera Rebutia and
Aylostera and of the subgenera within this last genus:
Aylostera and Mediolobivia. Finally we report the list of
taxa in our opinion belonging to genera Rebutia K.
Schum., and Aylostera Speg. About Rebutia K. Schum.,
we did not cite the infraspecific taxa since further
systematic investigations are still necessary. We only
cited R. wessneriana subsp. berilloydes since our sample
was identified as such and we preferred to maintain this
name for the molecular analysis. The same can be said
about R. einsteinii (Aylostera subgen. Mediolobivia in our
treatment). In any case we preferred not to accept the
combination Rebutia einsteinii Frič subsp. aureiflora
(Boed.) Hjertson (Hunt 2006), since it was not supported
by any morphological or biomolecular data. For this
reason the two taxa (R. einsteinii e R. aureiflora) should
be kept separated at the species level (until further data is
available).
In this treatment we propose necessary combinations
following the separation of Aylostera as a genus
independent from Rebutia. It is to remark that those
species belonging to subgen. Mediolobivia described as
representatives of genus Lobivia Britton & Rose by
Rausch (1986), were recombined as Rebutia by Sida
(1997) and Mosti (1999; 2000a; 200b).
With respect to the revision of Rebutia sect.
Digitorebutia (Mosti 1999; 2000a; 2000b), here inserted
in subgen. Mediolobivia, we added seven taxa: Rebutia
froehlichiana Rausch, R. iridescens F. Ritter, R.
tropaeolipicta F. Ritter, and the recently described R.
marieae La. Fisch & Halda, R. odehnalii Halda, Seda &
Sorma, R. raffaellii Mosti & Papini, R. rovidana Mosti &
Papini. In the same subgenus we added also the
Cylindrorebutia and Setirebutia: R. einsteinii Frič ex
Kreuz. & Buining, R. gonjianii Kiesling, R. aureiflora
Backeb., R. brighignae Mosti & Papini and R. fischeriana
Slaba. All these names are treated exhaustively
underneath and, where necessary, recombined.
In Hunt's (2006) checklist, only three species (R.
fabrisii, R. minuscula and R. padcayensis) of Rebutia K.
Schum. and only a few taxa (10) of Aylostera Speg., were
considered valid at the species level (all under Rebutia).
STEFANO MOSTI ET AL.,
2776
Among the ten species of Aylostera, six (with five
subspecies) were assigned to subgen. Aylostera, four
(with three subspecies) to subgen. Mediolobivia. A high
number of other taxa were synonymized with these taxa.
Moreover our studies in the field showed that considering
the species of the group Aylostera/Rebutia (particularly
those belonging to subgen. Mediolobivia) as few species
with wide geographical distribution range would be
distant from the real situation. We observed in the field
that these taxa are often composed by micropopulations
often living in rock fissures and separated from other
populations by long distances and geographical barriers.
However, the list of taxa belonging to genus
Aylostera reported as follows should not be considered
always as an assertion of the systematic validity of the
taxa, but in some cases only from a nomenclature point of
view. As a matter of fact, the number of “good species” of
subgen. Aylostera is probably lower than the total number
of names here reported, as stated, for instance in Hunt's
(2006) checklist. We assume that some of these taxa
should be probably synonymized in a revision of
Aylostera subgen. Aylostera. We recombined all the
available taxa under Aylostera, since an updated revision
of this subgenus is still lacking and attempts to
synonymize taxa without more exhaustive data may be
misleading.
For instance, our analysis of genetic distances (total
number of substitutions, see table 2) indicated that some
of the taxa considered synonyms by Hunt (2006) showed
a genetic distance comparable to that between taxa
considered different species by the same author. For
instance R. donaldiana is considered synonym of R.
fiebrigii by Hunt (2006), while the genetic distance
between these two species is 16, the same as the distance
between R. steinmannii and R. einsteinii, that are
considered “good” species in Hunt (2006) treatment.
For the above mentioned reason, in our opinion the
definitive number of species belonging to Genus
Aylostera cannot be yet established, even because of
different opinions about this aspect (Mosti, 1999; 2000a;
2000b and Hunt, 2006) and without a complete systematic
investigations.
Analytical key
1a. Axils of the floral scales with hairs, hairs and bristles or woolly ........................................................... 2 Aylostera
1b. Axils of the floral scales naked (occasionally with slight tomentum) ………………………….…………………..3
2a. Style and filaments coalescent with the internal side of the receptacular wall for its whole length; axils of the
floral scales with both hairs and bristles …………………………………................. Aylostera subgen. Aylostera
2b. Style and filaments free from the receptacular wall or coalescent with the internal side of receptacular wall only
for its lower part; axils of the floral scales with hairs or woolly but never bristly ………………………………....
…………………………………………………………………………………… Aylostera subgen. Mediolobivia
3a. Areoles inserted, more superficially, on the central (distal) part of the tubercle; scales of the floral tube narrow
and not imbricated. Spines quite fragile and subtle ...................................................................................... Rebutia
3b. Areoles on the tubercle in eccentric position, sunken in its higher (adaxial) part; floral tube scales broad and
imbricated. Spines more robust and thick. (Spines and floral tube lacked in Cintia) .................................................
……………………………………………………………………….. Weingartia (with Sulcorebutia and Cintia)
Rebutia K. Schum., Monatsschr. Kakteek. 5: 102. 1985.
Wessner). Habitat: environments mostly shady.
Small cactus (2-5 cm tall and 2-8 cm wide), flattenedglobular to globular or exceptionally sub-cylindrical.
Fusiform or rapiform root system. Spiralled or vertically
directed ribs scarcely developed, and resolved in low
isodiametrical, hexagonal tubercles spiralled or vertical.
Areoles small, round or oval or narrow elongated in the
centre of the tubercles , felted and initially without spines.
Spines often subtle and fragile, short to relatively short
(0.1-3 cm), bristly to acicular, adpressed or porrect, never
hooked, laterals and centrals often indistinguishable (15 to
35 in number), weak. Flowers (1-6.5 cm long and 1-5.5
cm wide), funnelform, arising only from a base or sides of
plant, never from the young areoles in the crown;
anthesis-time usually 2 days; stile and filaments not
coalescent to partially coalescent with the receptacle wall;
receptacular scales usually narrow, exceptionally broad;
scale axils of receptacle naked (occasionally with slight
tomentum). Fruit a small berry with persistent floral
remains, globose, naked, becoming papery at maturity.
Seeds black, conical cap shaped, with cuticula separated
from the testa and with verrucose testa and prominent
white hilum. Plants mostly self-fertile, but also sometime
self-sterile (Rebutia calliantha Bewer., R. kariusiana
Type: Rebutia minuscula K. Schum.
Checklist of species and infraspecific taxa
1. Rebutia calliantha Bewer., Sukkulentenkunde 2: 25.
1948. [= Rebutia minuscula subsp. wessneriana in
Muruaga et al., (2008)]
2. Rebutia fabrisii Rausch, Kakt., and. Sukk. 28(3): 52.
1977.
Rebutia kariusiana Wessner, Kakt. And. Sukk. XIV. 149.
1963.
4. Rebutia krainziana Kesselr., Succulentenkunde 2: 23.
1948.
5. Rebutia margarethae Rausch, Kakt. And. Sukk. 23(1):
4. 1972.
6. Rebutia marsoneri Werderm., Kakteenkunde 1937: 2.
1937.
7. Rebutia minuscula K. Schum., Monatsschr. Kakteenk.
NEW COMBINATIONS IN AYLOSTERA
5: 102. 1895. [Rebutia minuscula subsp. minuscula in
Muruaga et al., (2008) ].
8. Rebutia padcayensis Rausch, Kakt. & Sukkulent. XXI.
65. 1970.
2777
and filaments usually coalescent with the internal side of
the receptacular wall for the whole length of receptacle;
scale axils with both hairs and bristles. Fruits bristly.
Plants mostly self-fertile.
Type: Echinopsis pseudominuscula Speg.
9. Rebutia senilis Backeb., Kakteenfreund 1: 124. 1932.
[= R. minuscula subsp. minuscula in Muruaga et al.,
(2008)].
10. Rebutia violaciflora Backeb., Blatt. Kakteenf. 1935,
Pt 8. 1935. [Rebutia minuscula subsp. minuscula in
Muruaga et al., (2008)].
11. Rebutia wessneriana Bewer., Sukkulentenkunde 2:
24. 1948. [= Rebutia minuscula subsp. wessneriana in
Muruaga et al., (2008)].
11a. Rebutia wessneriana Bewer. subsp. berylloides
(Buining & Donald) Donald, Ashingtonia 2 (4): 71. 1976.
[= Rebutia minuscula in Hunt (2006)].
12. Rebutia xanthocarpa Backeb., Kakteenfreund 1: 131.
1932. [= R. minuscula subsp. minuscula in Muruaga et al.,
(2008)]
Aylostera Speg., Anal. Soc. Cient. Argent. 96: 75. 1923.
Small cactus (2-8 tall and 1-5 cm wide) flattenedglobular to cylindrical with rapiform or fusiform rootsystem. Ribs poorly to well developed forming spiralled
or vertical rows of more or less small, isodiametrical,
circular or hexagonal tubercles, sometime prominent.
Areoles small, round or oval or narrow elongated in centre
of the tubercles, felted and initially without spines. Spines
relatively short: (0.1-) 0.2-2 (-4) cm long, relatively weak
to always straight, bristly to acicular, pectinate or porrect,
never hooked, laterals and centrals often distinguishable
for a total amount of (4-) 7 to 40 (-50) ca. in number.
Flowers (0.8-5.5 cm long and 1.2-6 cm wide) funnelform
to campanulate with an almost cylindrical to cylindrical
receptacle, arising only from basal or lateral areoles, never
from the young areoles in the crown; anthesis-time
usually 2 days; tube scales usually narrow, exceptionally
broad; scale axils of receptacle with hairs, hairs and
bristles or woolly; style and filaments free from the
internal side of the receptacle wall or coalescent with the
receptacle wall for it whole length or only with the lower
part. Fruit a small berry with persistent floral remains,
flattened to roundish, naked, hairy or bristly, becoming
papery at maturity. Seeds brown to black, sometimes with
separated cuticula and with verrucose testa. Plants either
self-sterile or self-fertile.
Habitat: environments mostly exposed to open sun.
Type: Echinopsis pseudominuscula Speg.
Aylostera Speg. subgen. Aylostera
Plants flattened-globular, globular, sub-cylindrical, or
cylindrical; sometime with bronzed epidermis. Ribs
poorly developed from circular or hexagonal tubercles.
Spines up to 4.0 cm long and about 50 in number. Flowers
funnelform with, usually, a narrow cylindrical tube; style
Checklist of species and infraspecific taxa, including
taxonomic novelties
1. Aylostera albiareolata (F. Ritter) Mosti & Papini,
comb. nov.
Basionym: Rebutia albiareolata F. Ritter, Kakt. And.
Sukk. 28(4): 78. 1977.
Type: Bolivia. Tarija, Arce, "Padcaya", -/1/1963, Ritter
761 (holotype, U; isotype, ZSS-SR14416 (Eggli et al.,
1995)).
2. Aylostera albiflora (F. Ritter & Buining) Backeb.,
Descr. Cact. Nov. 3: 5. 1963.
3. Aylostera albipilosa (F. Ritter) Backeb., Descr. Cact.
Nov. 3: 5. 1963.
4. Aylostera albopectinata (Raush) Mosti & Papini,
comb. nov.
Basionym: Rebutia albopectinata Rausch, Kakt., and.
Sukk. 23 (9): 236. 1972.
Type: Bolivia. Sud Cinti, near Culpina, 3400 m, s.d.,
Rausch 312 (holotype, ZSS).
5. Aylostera azurduyensis J. de Vries, Succulenta 85(2):
54, 56, 59. 2006.
6. Aylostera brunescens (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia brunescens Rausch, Kakt. And Sukk.
23(9): 235. 1972.
Type: Bolivia. Chuquisaca, near Tarabuco, 3550 m, s.d.,
Rausch 480a (holotype, W; isotype ZSS).
7. Aylostera buiningiana (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia buiningiana Rausch, Kakt. And. Sukk.
23(4): 98. 1972.
Type: Argentina. Jujuy, near Iruya, 2700 m, s.d., Rausch
511 (holotype, W; isotype, ZSS).
8. Aylostera cajasensis (F. Ritter) Mosti & Papini, comb.
nov.
Basionym: Rebutia cajasensis F. Ritter, Succulenta 56(3):
64. 1977.
Type: Bolivia. Tarija, Mendez, near Cajas, 1963, Ritter
1141 (holotype, U; isotype, SGO-124626).
9. Aylostera cintiensis (F. Ritter) Mosti & Papini, comb.
nov.
Basionym: Rebutia cintiensis F. Ritter, Ashingtonia 2(10):
206. 1977:
2778
Type: Bolivia. Chuquisaca, Nor Cinti to the north of
Camargo, s.d., Ritter 938 (holotype, U; isotype, SGO124605).
10. Aylostera deminuta Britton & Rose, Cactaceae
(Britton & Rose) 4: 285. 1923.
11. Aylostera donaldiana (A.B.Lau & G.D.Rowley)
Mosti & Papini, comb. nov.
Basionym: Rebutia donaldiana A.B. Lau & G.D. Rowley,
Ashingtonia 1(7): 76. 1974.
Type: Bolivia. Santa Cruz, Valle Grande, Pucara, 2400
m, "growing in crevices in steep granite rocks facing
north", s.d., Lau 348 (holotype, HEID).
12. Aylostera fiebrigii (Gurke) Backeb., Kaktus-ABC:
274. 1936.
13. Aylostera flavistyla (F. Ritter) Mosti & Papini, comb.
nov.
Basionym: Rebutia flavistyla F. Ritter, Ashingtonia 3(1)
12. 1978.
Type: Bolivia. Tarija, Méndez, below Cajas, 1958, Ritter
756 (holotype, U; isotypes, SGO-124584, ZSSR14408).
14. Aylostera fulviseta (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia fulviseta Rausch, Kakt. & Sukkulent.
XXI. 29. 1970.
Type: Bolivia. Arque, near Padcaya, 2200 m, s.d., Rausch
319 (holotype, W; isotype ZSS).
15. Aylostera fusca (F. Ritter) Mosti & Papini, comb.
nov.
Basionym: Rebutia fusca F. Ritter, Kakt. And. Sukk.
28(4): 78 1977.
Type: Bolivia. Tarija, "westlich Tarija", 1959, Ritter 940
(holotype, U).
16. Aylostera heliosa (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia heliosa Rausch, Kakt. And. Sukk.
XXI. 30. 1970.
Type: Bolivia. Tarija, near Tarija on the road to Narvaez,
2400-2500 m, s.d., Rausch 314 (holotype, W; isotype,
ZSS).
16a. Aylostera heliosa (Rausch) Mosti & Papini subsp.
cajasensis (Donald) Mosti & Papini, comb. nov. & stat.
nov.
Basionym: Rebutia heliosa Rausch var. cajasensis
Donald, Aschingtonia 3(5-6): 144-145. 1979.
Type: Bolivia. Tarija: Abra Cajas, 2400 m, s. d., Lau 405
(holotype, K).
16b. Aylostera heliosa (Rausch) Mosti & Papini subsp.
condorensis (Donald) Mosti & Papini, comb. nov. & stat.
nov.
STEFANO MOSTI ET AL.,
Basionym: Rebutia heliosa Rausch var. condorensis
Donald, Aschingtonia 3(5-6): 143-144. 1979.
Type: Bolivia. Tarija: Abra Condor, 2500 m, s. d., Lau
401 (holotype, K).
17. Aylostera hoffmannii (Diers & Rausch) Mosti &
Papini, comb. nov.
Basionym: Rebutia hoffmannii Diers & Rausch, Kakt.
And. Sukk. 28(5): 105. 1977.
Type: Argentina. Salta, near Santa Victoria, s.d., Rausch
521a (holotype, ZSS).
18. Aylostera jujuyana (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia jujuyana Rausch, Kakt. And. Sukk.
24(7): 147-148. 1973.
Type: Argentina. Jujuy, Quebrada de Humahuaca, s.d.,
Rausch 220 (holotype, ZSS). .
19. Aylostera kieslingii (Rausch) Mosti & Papini, comb.
nov.
Basioym: Rebutia kieslingii Rausch, Kakt. And. Sukk.
28(8): 177-178. 1977.
Type: Argentina. Salta, near Caspala, 3200 m, s.d.
Rausch 694 (holotype, ZSS).
20. Aylostera kupperiana (Boed.) Backeb. in Backeb. &
F.M. Knuth, Kaktus-ABC 275. 1936.
21. Aylostera mamillosa (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia mamillosa Rausch, Succulenta 51(4):
69. 1972.
Type: Bolivia. Chuquisaca, Sud Cinti, west of Camargo,
3300 m, s. d., Rausch 302 (holotype, ZSS).
22. Aylostera mandingaensis R. Wahl & Jucker, Kakteen
Sukk. 59(4): 101-105. 2008.
23. Aylostera muscula (F. Ritter & Thiele) Backeb.,
Descr. Cact. Nov.: 3. 5. 1963.
24. Aylostera narvacense Cárdenas, Cact. Succ. J. 43(6):
245. 1971.
25. Aylostera nitida (F. Ritter) Mosti & Papini, comb.
nov.
Basionym: Rebutia nitida F. Ritter, Ashingtonia 3(1): 14.
1978.
Type: Bolivia. Tarija, Mendez, Cajas, 2000 m, 1958,
Ritter 769 (holotype, U [not found]; isotypes, ZSSS10437 [seeds only], SGO-124596).
26. Aylostera nogalensis (F. Ritter) Mosti & Papini,
comb. nov.
Basionym: Rebutia nogalensis F. Ritter, Kakt. And. Sukk.
28(4): 78. 1977.
NEW COMBINATIONS IN AYLOSTERA
Type: Bolivia. Chuquisaca, Azurduy, "Tarvita, rocky
slopes", 1958, Ritter 768 (holotype, U; isotype, SGO124595).
27. Aylostera patericalyx (F. Ritter) Mosti & Papini,
comb. nov.
Basionym: Rebutia patericalyx F. Ritter, Kakt. And.
Sukk. 28(4): 78. 1977.
Type: Bolivia. Chuquisaca, Sud Cinti, "La Cueva", 1963,
Ritter 757 (holotype, U; isotypes, SGO-124585, ZSSSR14409).
28. Aylostera perplexa (Donald) Mosti & Papini, comb.
nov.
Basionym: Rebutia perplexa Donald, Ashingtonia 3(5-6):
150. 1980.
Type: Bolivia.Tarija/Chuquisaca, prope Rio Pilaya, s.d.,
Lau 329a (holotype, K; isotype, ZSS).
29. Aylostera pseudodeminuta (Backeb.) Backeb.,
Backeb. & F.M. Knuth, Kaktus-ABC: 275. 1936.
30. Aylostera pseudominuscula (Speg.) Speg., Anal. Soc.
Ci. Argent. 96: 75. 1923.
31. Aylostera pulchella (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia pulchella Rausch, Kakt. And. Sukk.
23(12): 340. 1972.
Type: Bolivia. Chuquisaca, to the north of Padilla, 2200
m, s.d., Rausch 320 (holotype, W; isotype, ZSS).
32. Aylostera pulvinosa (F. Ritter & Buining) Backeb.,
Descr. Cact. Nov. 3: 5. 1963.
33. Aylostera robustispina (F. Ritter) Mosti & Papini,
comb. nov.
Basionym: Rebutia robustispina F. Ritter, Succulenta
56(3): 64. 1977.
Type: Bolivia. Tarija, mountains N of Tarija, 1966, Ritter
763 (holotype, U; isotypes, ZSS-SR14421, ZSSSR14422).
34. Aylostera rubiginosa (F. Ritter) Backeb., Descr. Cact.
Nov. 3: 5. 1963.
35. Aylostera schatzliana (Rausch) Mosti & Papini,
comb. nov.
Basionym: Rebutia schatzliana Rausch, Kakt. And. Sukk.
26(11): 224-225. 1975.
Type: Bolivia. Chuquisaca, Nor Cinti, near Pucara, 3200
m, s.d., Rausch 640 (holotype, ZSS).
36. Aylostera simoniana (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia simoniana Rausch, Kakt. And. Sukk.
35(9): 205. 1984.
Type: Bolivia. Chuquisaca, Sud Cinti, south of and above
La Cueva, 3500 m, s.d., Rausch 739 (holotype, ZSS).
2779
37. Aylostera spegazziniana (Backeb.) Backeb., Backeb.
& F.M Knuth, Kaktus-ABC 276. 1936.
38. Aylostera spinosissima (Backeb.) Backeb., Backeb. &
F.M Knuth, Kaktus-ABC 275. 1936.
39. Aylostera sumayana (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia sumayana Rausch, Succulenta 65(4):
74. 1986.
Type: Bolivia. Chuquisaca, Sud Cinti, near Sumaya, 3200
m, s.d., Rausch 738 (holotype, ZSS).
40. Aylostera supthutiana (Rausch) Mosti & Papini,
comb. nov.
Basionym: Rebutia supthutiana Rausch, Kakt. And. Sukk.
27(6): 121-122. 1976.
Type: Bolivia. Tarija, O'Connor, at Tambo, s.d., Rausch
629 (holotype, ZSS).
41. Aylostera tamboensis (F. Ritter) Mosti & Papini,
comb. nov.
Basionym: Rebutia tamboensis F. Ritter, Ashingtonia
2(10): 207. 1977.
Type: Bolivia. Tarija, O'Connor, east of Tarija, "at the
upper end of the Tambo gorge", 2500 m, 1962, Ritter
1142 (holotype, U; isotype, SGO-124627). i
42. Aylostera tarijensis (Rausch) Mosti & Papini, comb.
nov.
Basionyn: Rebutia tarijensis Rausch, Kakt. and. Sukk.
26(9): 195-196. 1975.
Type: Bolivia. Tarija, east of Tarija, 2600 m, s.d., Rausch
87 (holotype, ZSS).
43. Aylostera tarvitaensis (F. Ritter) Mosti & Papini,
comb. nov.
Basionym: Rebutia tarvitaensis F. Ritter, Kakt. and. Sukk.
28(4): 78. 1977.
Type: Bolivia. Chuquisaca, Azurduy, at Tarvita, 1958,
Ritter 773 (holotype, U; isotypes, SGO-124599, ZSSSR14432).
44. Aylostera tuberosa (F. Ritter) Backeb., Descr. Cact.
Nov. 3, 5. 1963.
45. Aylostera vallegrandensis (Cárdenas) Mosti & Papini,
comb. nov.
Basionym: Rebutia vallegrandensis Cárdenas, Cact. Succ.
J. (Los Angeles) 42: 35. 1970.
Type: Bolivia. Santa Cruz, Valle Grande, near Candelaria
on the way to Rio Piraymiri, 2000 m, s.d., Card. 6307
(holotype, LIL (Cárdenas Herbarium)).
46. Aylostera vulpina (F. Ritter) Mosti & Papini, comb.
nov.
Basionym: Rebutia vulpina F. Ritter, Succulenta 56(3):
66. 1977.
STEFANO MOSTI ET AL.,
2780
Type: Bolivia. Tarija, Méndez, west of Tarija, 1959,
Ritter 939 (holotype, U; isotype, SGO-124606). .
47. Aylostera wahliana (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia wahaliana Rausch, Succulenta 64(12):
257. 1985.
Type: Bolivia. Tarija, at Cuesta de Sama, s.d., Rausch
645 (holotype, ZSS).
48. Aylostera walteri (Diers) Mosti & Papini, comb. nov.
Basionym: Rebutia walteri Diers, Kakt. And. Sukk. 40(8):
189-190. 1989.
Type: Argentina, Salta, Santa Victoria, 2500-2700 m,
s.d., Hoffmann 1960a (holotype, KOELN).
49. Aylostera waltheriana (Backeb.) Y. Ito, Explan.
Diagr. Austroechinocactinae: 129. 1957.
Aylostera Speg. subgen. Mediolobivia
Rowley, Cactus World 27(2): 88-90. 2009
(Backeb.)
Plants flattened-globular, globular, sub-cylindrical or
cylindrical; often with bronzed epidermis. Ribs well or
sometimes poorly developed forming spiralled or vertical
rows of rounded tubercles. Spines often with thick
bulbous bases up to 1-20 (-60 mm in Aylostera aureiflora)
mm long and 4-20 in number. Flowers funnelform or
campanulate; style and filaments usually coalescent with
the internal side of the receptacular wall for the lower part
of the receptacle, or free from the receptacular wall; scale
axils pilose (to very woolly) but never bristly. Fruits
usually naked or only weakly pilose. Plants either selfsterile or self-fertile.
53. Aylostera aureiflora (Backeb.) Mosti & Papini, comb.
nov.
Basionym: Rebutia aureiflora Backeb., Kakteenfreund 1:
124. 1932.
Type: Argentina. Salta: Quebrada del Toro, s. d., s. coll.
(lectotype (Hjertson, 2003), photograph in KakteenFreund 1: 24. 1932).
54. Aylostera brachyantha (Wessner) Mosti & Papini,
comb. nov.
Basionym: Lobivia brachyantha Wessner, Kakt. And.
Sukk. 1937 (9): 129, 207. 1937.
Type: Bolivia. Potosi, near Potosi, s.d., s.n. (lectotype
(Hunt & Taylor, 2006), illustration in Kakt. And. Sukk.
1937 (9): 131-132.).
55. Aylostera brunneoradicata (F. Ritter) Mosti & Papini,
comb. nov.
Basionym: Rebutia brunneoradicata F. Ritter, Kakt. And.
Sukk. 28(4): 77-78. 1977.
Type: Bolivia. Tarija, Mendez, "San Antonio", 1962,
Ritter 1109 (holotype, U).
56. Aylostera brighignae (Mosti & Papini) Mosti &
Papini, comb. nov.
Basionym: Rebutia brighignae Mosti & Papini, Cactus &
Co. 9(1): 60-61. 2005.
Type: Argentina. Jujuy, Abra de Pives, s.d., Rausch 751,
cult. S. Mosti (holotype, FT!).
Type: Rebutia haagei Frič & Shelle.
57. Aylostera camargoensis (Rausch) Mosti & Papini,
comb. nov.
Basionym: Rebutia camargoensis Rausch, Succulenta
55(3): 42. 1976.
Checklist of species and infraspecific taxa, including
taxonomic novelties
Type: Bolivia. Chuquisaca, Nor Cinti, west of Camargo,
3200 m, s.d., Rausch 311 (holotype, ZSS)..
50. Aylostera amblypetala (F. Ritter) Mosti & Papini,
comb. nov.
Basionym: Rebutia rosalbiflora F. Ritter var. amblypetala
F. Ritter, Kakt. And. Sukk. 28(4): 76. 1977.
58. Aylostera canacruzensis (Rausch) Mosti & Papini,
comb. nov.
Basionym: Rebutia canacruzensis Rausch, Kakt. And.
Sukk. 27(3): 49-50. 1976.
Type: Bolivia. Potosi, Sud Chichas, Impora above Tarata,
s.d., Ritter 1119 (holotype, U [not found]). i
Type: Bolivia. Chuquisaca, Nor Cinti, near Cana Cruz
(Caña Cruz?), 3700 m, s.d., Rausch 642 (holotype, ZSS).
51. Aylostera applanata (Rausch) Mosti & Papini, comb.
nov.
Basionym: Lobivia steinmannii (Solms-Laub.) Backebg.
var. applanata Rausch, Lobivia 85: 145, 129. 1986.
59. Aylostera carmeniana (Rausch) Mosti & Papini,
comb. nov.
Basionym: Rebutia carmeniana Rausch, Kakt. And. Sukk.
29(5): 105. 1978.
Type: Bolivia. Chuquisaca, north of Camargo, Rio
Hondo, 3600 m, s.d., Rausch 486 (holotype, ZSS).
Type: Argentina. Salta, "in ravines near Caspala", 3100
m, s.d., Rausch 690 (holotype, ZSS).
52. Aylostera atrovirens (Backeb.) Mosti & Papini, comb.
nov.
Basionym: Lobivia atrovirens Backeb. in Backeb. & F.M.
Knuth, Kaktus-ABC, 242, 414. 1936. Locality at the
protologue: Argentina. Salta to Jujuy, from high
mountains (to be typified).
60. Aylostera christinae (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia christinae Rausch, Kakt. And. Sukk.
26(7): 145. 1975.
NEW COMBINATIONS IN AYLOSTERA
Type: Argentina. Salta, between Nazareno and Rodeo,
3500 m, s.d., Rausch 492a (holotype, W).
61. Aylostera cincinnata (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia cincinnata Rausch, Kakt. And. Sukk.
27(1): 4. 1976.
Type: Bolivia. Potosi, near Cuchu Ingenio, 3600 m, s.d.,
Rausch 300 (holotype, ZSS).
62. Aylostera colorea (F. Ritter) Mosti & Papini, comb.
nov.
Basionym: Rebutia colorea F. Ritter, Kakt. And. Sukk.
28(4): 78. 1977.
Type: Bolivia. Tarija, Mendez, "San Antonio", 1962,
Ritter 1106 (holotype, U).
63. Aylostera crassa (Rausch) Mosti & Papini, comb.
nov.
Basionym: Lobivia haagei (Fric & Shelle) Wessner var.
crassa Rausch, Lobivia 85: 140, 56. 1986.
Type: Bolivia. Tarija, "in regionibus altis apud Sama", at
Iscayachi, s.d., Rausch 501 (holotype, ZSS).
64. Aylostera diersiana (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia diersiana Rausch, Kakt. And. Sukk.
26(2): 25-26. 1975.
Type: Bolivia. Chuquisaca, Sud Cinti, Yuquina near
Culpina, 3200 m, s.d., Rausch 631 (holotype, ZSS).
64a. Aylostera diersiana (Rausch) Mosti & Papini subsp.
atrovirens (Rausch) Mosti & Papini, comb. nov. & stat.
nov.
Basionym: Rebutia diersiana Rausch var. atrovirens
Rausch, Kakt. And. Sukk. 26(2): 26.
Type: Bolivia. Chuquisaca: Sud Cinti, above Salitre near
Culpina, 3200 m, s. d., Rausch 633 (holotype, ZSS).
65. Aylostera einsteinii (Frič ex Kreuz. & Buining) Mosti
& Papini, comb. nov.
Basionym: Rebutia einsteinii Friè ex Kreuz. & Buining,
Succulenta 1949: 65. 1949.
Type: Argentina. Jujuy, Quebrada del Toro, s.d., Lau
477? (lectotype (Hjertson, 2003), photogr. in Kaktusar 3:
16. 1932)
2781
Type: Argentina. Jujuy, s.d., L. Fischer 54 (holotype, PR).
68. Aylostera friedrichiana (Rausch) Mosti & Papini,
comb. nov.
Basionym: Rebutia friedrichiana Rausch, Succulenta
55(6): 103. 1976.
Type: Bolivia. Chuquisaca, Sud Cinti, near La Cueva,
3500 m, s.d., Rausch 646 (holotype, ZSS).
69. Aylostera froehlichiana (Rausch) Mosti & Papini,
comb. nov.
Basionym: Rebutia froehlichiana Rausch, Succulenta
54(12): 226. 1975.
Type: Bolivia. Chuquisaca, Sud Cinti, near Yuquina,
3200 m, s.d., Rausch 649 (holotype, ZSS).
70. Aylostera gavazzii (Mosti) Mosti & Papini, comb.
nov.
Basionym: Rebutia gavazzii Mosti, Cactus & Co. 3(4):
207-209. 1999.
Type: Bolivia, Tarija, s.d., Rausch 493, cult. S. Mosti
(holotype, FI!).
71. Aylostera gonjianii (Kiesling) Mosti & Papini, comb.
nov.
Basionym: Rebutia gonjianii Kiesling, Bol. Soc. Arg. Bot.
15: 132. 1973.
Type: Argentina. Jijuy, Tilcara, "a cuatro horas a mulas
oeste de Huacalera", Jan. 1973, B. Gonjian 2 (holotype,
LP).
72. Aylostera haagei (Frič & Shelle) Mosti & Papini,
comb. nov.
Basionym: Rebutia haagei Frič & Shelle, Kaktusar 1: 88.
1930.
Type: Argentina, Jujuy, s.d., s.n. (lectotype (Hjertson,
1994), photograph in Frič & Schelle. 1930).
72a. Aylostera haagei (Frič & Shelle) Mosti & Papini
subsp. elegantula (Rausch) Mosti & Papini, comb. nov.
Basionym: Lobivia haagei Frič & Shelle var. elegantula
Rausch, Lobivia 85: 140, 57. 1986.
Type: Argentina. Jujuy, Río San Juan de Oro, s.d.,
Rausch 502 (holotype, ZSS).
66. Aylostera eos (Rausch) Mosti & Papini, comb. nov.
Basionym: Rebutia eos Rausch, Succulenta 51(1):2. 1972.
72b. Aylostera haagei (Fric & Schelle) Mosti & Papini
subsp. mudanensis (Rausch) Mosti & Papini, comb. nov.
& stat. nov.
Basionym: Rebutia mudanensis Rausch, Kakt. And. Sukk.
27(8): 169. 1976.
Type: Argentina. Jujuy, near Tafna, 3600 m, s.d., Rausch
333 (holotype, W).
Type: Argentina. Salta: east of Mudana near Santa Ana,
3500-4300 m, s. d., Rausch 689 (holotype, ZSS).
67. Aylostera fischeriana (Slaba) Mosti & Papini, comb.
nov.
Basionym: Rebutia fischeriana Slaba in Kaktusy (Brno)
38(3): 69. 2002.
73. Aylostera haefneriana (Cullmann) Mosti & Papini,
comb. nov.
Basionym: Mediolobivia haefneriana Cullmann, Kakt.
And. Sukk. 6(2): 119. 1955. Locality at the protologue:
Bolivia. Potosi, s.d., s.n. (to be typified).
2782
74. Aylostera huasiensis (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia huasiensis Rausch, Kakt. And. Sukk.
28(2): 25. 1977.
Type: Bolivia. Chuquisaca, Inca Huasi, near Culpina,
3300 m, s.d., Rausch 313 (holotype, ZSS).
75. Aylostera iridescens (F. Ritter) Mosti & Papini, comb.
nov.
Basionym: Rebutia iridescens F. Ritter, Kakt. And. Sukk.
28(4): 76. 1977.
Type: Bolivia. Potosi: Sud Chicas, W of Mal Paso, /-//1962, Ritter 1434 (holotype, U).
76. Aylostera iscayachensis (Rausch) Mosti & Papini,
comb. nov.
Basionym: Rebutia ischayachensis Rausch, Succulenta
56(1): 3. 1977.
Type: Bolivia. Tarija, Mendez, above Cana Cruz, near
Iscayachi, 3500 m, s.d., Rausch 335b (holotype, ZSS).
77. Aylostera knizei (Rausch) Mosti & Papini, comb. nov.
Basionym: Lobivia pygmaea (R. E. Fr.) Backeb. var.
knizei Rausch, Lobivia 85: 144, 116. 1986.
Type: Bolivia. Oruro, prope Challapata, 3800 m, s.d.,
Rausch 676a (holotype, ZSS).
78. Aylostera leucacantha (Rausch) Mosti & Papini,
comb. nov.
Basionym: Lobivia steinmannii (Solms-Laub.) Backebg.
var. leucacantha Rausch, Lobivia 85: 145, 129. 1986.
STEFANO MOSTI ET AL.,
Basionym: Lobivia steinmannii (Solms-Laub.) Backebg.
var. melanocentra Rausch, Lobivia 85: 146, 129. 1986).
Type: Bolivia. Tarija, ab Iscayachi, "ab occidentem
versus", 3600 m, s.d., Rausch 744 (holotype, ZSS).
83. Aylostera minor (Rausch) Mosti & Papini, comb. nov.
Basionym: Rebutia diersiana Rausch var. minor Rausch,
Succulenta 58(11): 258. 1979.
Type: Bolivia. Sud Cinti at Yuquina, near Culpina, 3600
m, s.d., Rausch 630 (holotype, ZSS).
84. Aylostera mixticolor (F. Ritter) Mosti & Papini,
comb. nov.
Basionym: Rebutia mixticolor F. Ritter, Kakt. And. Sukk.
28(4): 77. 1977.
Type: Bolivia. Tarija, Méndez, "San Antonio", 1966,
Ritter 1108 (holotype, U).
85. Aylostera nazarenoensis (Rausch) Mosti & Papini,
comb. nov.
Basionyon:
Digitorebutia
nazarenoensis
Rausch,
Succulenta: 58(8): 186. 1979.
Type: Argentina. Salta, near Nazareno, 3350 m, s.d.,
Rausch 484 (holotype, ZSS).
86. Aylostera nigricans (Wessner) Mosti & Papini, comb.
nov.
Basionym:
Lobivia
nigricans
Wessner,
Beitr.
Sukkulentenk. Sukkulentenpflege 3: 51. 1940.
Type: Bolivia. Chuquisaca, Cinti australis, supra La
Cueva, 3700 m, s.d., Rausch 644 (holotype, ZSS).
Type: Argentina. Salta, s.d., s.n. (lectotype (Hjertson,
2005), photograph of flowering plant showing flower
from the side in Beitr. Sukkulentenk. Sukkulentenpflege
3: 51. 1940).
79. Aylostera leucanthema (Rausch) Mosti & Papini,
comb. nov.
Basionym: Rebutia leucanthema Rausch, Kakt. And.
Sukk. 26(6): 125. 1977.
86a. Aylostera nigricans (Wessner) Mosti & Papini
subsp. albispina (Rausch) Mosti & Papini, comb. nov.
Basionym: Lobivia nigricans Wessner var. albispina
Rausch, Lobivia 85: 142, 98, 100. 1986.
Type: Bolivia. Chuquisaca, Nor Cinti, Cana Cruz, 3600
m, s.d., Rausch 305 (holotype, ZSS).
Type: Argentina. Salta, Escoipe (Cuesta de Obispo), 2700
m, s.d. Rausch 771 (holotype, ZSS).
80. Aylostera major (Rausch) Mosti & Papini, comb. nov.
Basionym: Lobivia steinmannii (Solms-Laub.) Backebg.
var. major Rausch, Lobivia 85: 146, 129. 1986.
87. Aylostera oculata (Werderm.) Mosti & Papini, comb.
nov.
Basionym: Rebutia oculata Werderm., Kakt., and. Sukk.
25, tab 99. 1935.
Type: Argentina. Jujuy, apud Tafna, 3600 m, s.d., Rausch
334 (holotype, ZSS).
81. Aylostera marieae (Lad.Fisch. & Halda) Mosti &
Papini, comb. nov.
Basionym: Rebutia marieae Lad.Fisch. & Halda, Acta
Mus. Richnov., Sect. Nat. 9(1): 71-72. 2002.
Type: Argentina. Jujuy, 2001-1-15, L. Fischer s.n.
(holotype, PR).
82. Aylostera melanocentra (Rausch) Mosti & Papini,
comb. nov.
Type: Argentina, Jujuy, s.d., s.n. (lectotype, icon in Kakt.
And. Sukk. 25: t. 99. 1935).
Syn.: Lobivia euanthema Backeb., Backeb. & F. M.
Knuth, Kaktus-ABC, 240, 414. 1936.
87a. Aylostera oculata (Werderm.) Mosti & Papini
subsp. tilcarensis (Rausch) Mosti & Papini, comb. nov. &
stat. nov.
Basionym: Lobivia euanthema Backeb. var. tilcarensis
Rausch, Lobivia 85: 138, 36. 1986.
NEW COMBINATIONS IN AYLOSTERA
Type: Argentina. Jujuy: "in summis montibus apud
Tilcara", s. d., Rausch 700 (holotype, ZSS).
88. Aylostera odehnalii (Halda, Seda & Sorma) Mosti &
Papini, comb. nov.
Basionym: Rebutia odehnalii Halda, Seda & Sorma, Acta
Mus. Richnov., Sect. Nat. 9(1): 72-73. 2002.
Type: Bolivia. s.d., J.J. Halda, J. Odehnal & V. Seda, s.n.
(holotype, PR).
89. Aylostera orurensis (Backeb.) Mosti & Papini, comb.
nov.
Basionym: Lobivia orurensis Backeb., Backeb. & F.M.
Knuth, Kaktus-ABC: 243, 415. 1936. Locality at the
protologue: Bolivia. Oruro, Poopo, near Oruro, 3800 m.
(to be typified).
90. Aylostera pallida (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia pallida Rausch, Succulenta 56(10):
234. 1977.
Type: Bolivia. Chuquisaca, Sud Cinti, around La Cueva,
3500 m, s.d., Rausch 645 (holotype, ZSS).
91. Aylostera parvula (Rausch) Mosti & Papini, comb.
nov.
Basionym: Lobivia steinmannii (Solms-Laub.) Backebg.
var. parvula Rausch, Lobivia 85: 146, 129. 1986.
Type: Bolivia. Potosi, Cucho Ingenio, 3800 m, s.d.,
Rausch 296 (holotype, ZSS).
92. Aylostera pauciareolata (F. Ritter) Mosti & Papini,
comb. nov.
Basionym: Rebutia pauciareolata F. Ritter, Kakt. And.
Sukk. 28(4): 77. 1977.
Type: Bolivia. Tarija, Mendez, "San Antonio", 1962,
Ritter 1121 (holotype, U; isotype, SGO-124623).
93. Aylostera pelzliana (Rausch) Mosti & Papini, comb.
nov.
Basionym: Lobivia haagei (Frič & Schelle) Wessner var.
pelzliana Rausch, Lobivia 85: 140, 57. 1986.
Type: Argentina. Jujuy, apud Tafna, 3600 m, s.d., Rausch
333a (holotype, ZSS).
94. Aylostera polypetala (Rausch) Mosti & Papini, comb.
nov.
Basionym: Lobivia pygmaea (R.E. Fr.) Backeb. var.
polypetala Rausch, Lobivia 85: 144, 116. 1986.
Type: Bolivia. Potosi, Cucho Ingenio, 3700 m, s.d.,
Rausch 301 (holotype, ZSS).
95. Aylostera pseudoritteri (Raush) Mosti & Papini,
comb. nov.
Basionym: Lobivia atrovirens Backeb. var. pseudoritteri
Rausch, Lobivia 85: 137, 13. 1986.
Type: Bolivia. Tarija, Abra de Sama, s.d., Rausch 506
(holotype, ZSS).
2783
96. Aylostera pygmaea (R.E. Fr.) Mosti & Papini, comb.
nov.
Basionym: Echinopsis pygmaea R.E. Fr., Nova Acta
Regiae Soc. Upsal. ser. 4, 1(1): 120. 1905).
Type: Argentina. Prov. Jujuy, Yavi, c. 3400 m, s.d., R.E
Fries 999 (lectotype (Hjertson, 1994), S).
97. Aylostera raffaellii (Mosti & Papini) Mosti & Papini,
comb. nov.
Basionym: Rebutia raffaellii Mosti & Papini, Cactus &
Co. 9(1): 56-57. 2005.
Type: Argentina. Jujuy, presso Tafna, s.d., Rausch 333c,
cult. S. Mosti (holotype, FT!).
98. Aylostera raulii (Rausch) Mosti & Papini, comb. nov.
Basionym: Rebutia raulii Rausch, Kakt. And. Sukk.
31(6): 171. 1980.
Type: Bolivia. Chuquisaca, north of Camargo, at the Rio
Honda, s.d., Rausch 485 (holotype, ZSS).
99. Aylostera ritteri (Wessner) Mosti & Papini, comb.
nov.
Basionym: Lobivia ritteri Wessner, Beitr. Sukkulentenk.
Sukkulentenpflege 1: 3. 1938.
Type: Bolivia. Tarija: Iscayachi Ritter 03/-/1931 s. n.
(lectotype (Hjertson, 2005), illus. in Beitr. z. Sukk. u.pfl.1:3. 1938).
100. Aylostera rovidana (Mosti & Papini) Mosti &
Papini, comb. nov.
Basionym: Rebutia rovidana Mosti & Papini, Cactus &
Co. 9(1): 58-59. 2005.
Type: Argentina. Jujuy, presso Tafna, s.d., Rausch 333b,
cult. S. Mosti (holotype, FT!).
101. Aylostera steinmannii (Solms) Backeb., Cactaceae
(Backeberg) 3: 1528. 1959.
Basionym: Echinocactus steinmannii Solms, Bot. Zeit.
65(1): 133. 1907.
Type: Bolivia. s. d., s. coll. (lectotype (Hjertson, 2005),
fig. 4 in Bot. Zeit. 65(1): plate 2. 1907).
102. Aylostera tafnaensis (Rausch) Mosti & Papini,
comb. nov.
Basionym: Lobivia pygmaea (R. E. Fr.) Backeb. var
tafnaensis Rausch, Lobivia 85: 144, 116. 1986.
Type: Argentina. Jujuy, apud Tafna, 4000 m, s.d., Rausch
508a (holotype, ZSS).
103. Aylostera torquata (F. Ritter & Buining) Mosti &
Papini, comb. nov.
Basioym: Rebutia torquata F. Ritter & Buining,
Succulenta 56(3): 63. 1977.
Type: Bolivia, Potosi, Sud Chichas, "Mal Paso", /11/1962, Ritter 1117 (holotype, U).
2784
104. Aylostera tropaeolipicta (F. Ritter) Mosti & Papini,
comb. nov.
Basionym: Rebutia tropaeolipicta F. Ritter, Kakt. And.
Sukk. 28(4): 78. 1977.
Type: Bolivia. Potosi, Sud Chichas, "Mal Paso", 1962,
Ritter 1114 (holotype, U; isotype, ZSS).
105. Aylostera tuberculata (Rausch) Mosti & Papini,
comb. nov.
Basionym: Lobivia steinmannii (Solms-Laub.) Backebg.
var. tuberculata Rausch, Lobivia 85: 145, 129. 1986.
Type: Bolivia, Cinti meridionalis, in pago La Cueva, s.d.,
Rausch 743 (holotype, ZSS).
106. Aylostera violaceostaminata (Rausch) Mosti &
Papini, comb. nov.
Basionym:
Lobivia
pygmaea
R.E.
Fr.
var.
violaceostaminata Rausch, Lobivia 85: 144, 116. 1986.
Type: Bolivia. Chuquisaca, Cinti australis, apud La
Cueva, 3300 m, s.d., Rausch 742 (holotype, ZSS).
107. Aylostera violascens (F. Ritter) Mosti & Papini,
comb. nov.
Basionym: Rebutia violascens F. Ritter, Kakt. And. Sukk.
28(4): 76. 1977.
Type: Bolivia. Chuquisaca, Nor Cinti, north of Camargo,
1958, Ritter 352 (holotype, U).
108. Aylostera yuncharasensis (Rausch) Mosti & Papini,
comb. nov.
Basionym:
Lobivia
atrovirens
Backeb.
var.
yuncharasensis Rausch, Lobivia 85: 137, 13. 1986.
Type: Bolivia. Tarija, Yunchara, s.d., Rausch 91
(holotype, ZSS).
109. Aylostera yuquinensis (Rausch) Mosti & Papini,
comb. nov.
Basionym: Rebutia yuquinensis Rausch, Kakt. And. Sukk.
31(10): 307. 1980.
Type: Bolivia. Chuquisaca, Sud Cinti, Yuquina, 3200 m
(9800 ft), s.d., Rausch 632 (holotype, ZSS).
110. Aylostera zecheri (Rausch) Mosti & Papini, comb.
nov.
Basionym: Rebutia zecheri Rausch, Succulenta 56(2): 30.
1977.
Type: Bolivia. Tarija, Iscayachi, near Pueblo Viejo, 3000
m (9850 ft), s.d., Rausch 650 (holotype, ZSS).
References
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(Received for publication 20 April 2011)