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A synopsis of the genus Cleistocactus Lemaire (Cactaceae)
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DOI: 10.25223/brad.n34.2016.a6
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Bradleya 34/2016
pages 148–186
A synopsis of the genus Cleistocactus Lemaire (Cactaceae)

Martin Lowry
Northgate Avenue, Macclesfield, UK (email: m.lowry@hull.ac.uk).
Summary: The history of the genus Cleistocactus Universel allowing Lemaire to make an addition
Lem. (Cactaceae) is reviewed and significant eras to his preliminary description. This presumably
outlined. A conservative species concept has been accounts for the curious separation of the first
employed to group the over 300 published names description over two widely-separated pages of the
into 24 species and 3 heterotypic subspecies. A volume. Monville described the flowers as being
complete synonymy is given for each of the bright orange and very different in shape from
accepted species along with an expanded other cerei; more like those of Epiphyllum
description, distribution maps, ecological region truncatum (now Schlumbergera truncata, the
associations and a brief commentary. The Christmas cactus). It appears Lemaire did not see
descriptions are expanded from the respective the flowers himself and continued to call the plant
protologues to include additional data observed Cereus baumannii in subsequent publications
both in the field and in ex-situ culture. over several years. Only when eventually
considering the production of a monograph did he
Zusammenfassung: Die Geschichte der Gattung accept that the genus Cereus needed dividing.
Cleistocactus Lem. (Cactaceae) wird betrachtet Thus, in 1860, he created the genus Aporocactus
und bedeutende Perioden skizziert. Ein and transferred his Cereus baumannii there along
konservatives Artkonzept wurde angewendet, um with C. flagelliformis. This unfortunate mistake
die über 300 publizierten Namen in 24 Arten und was rapidly pointed out to him by a “distinguished
drei heterotypische Unterarten zu gruppieren. botanist” and by an amateur cactophile, neither
Eine komplette Synonymie wird für jede der of whom he names. The amateur botanist was
anerkannten Arten gegeben, zusammen mit einer very likely Schlumberger of Rouen who had
erweiterten Beschreibung, Verbreitungskarten, amassed a large collection often used by Lemaire.
Zuordnung zu ökologischen Regionen und einem Lemaire subsequently made a trip to the botanical
kurzen Kommentar. Die Beschreibungen werden garden at Caen to see the flowers of Cereus
ausgehend von den entsprechenden Protologen colubrinus Hort. which had been described as
um zusätzliche Beobachtungsdaten ergänzt, die identical to those of his C. baumannii.
sowohl in der Natur als auch in ex-situ-Kulturen Recognising his error he published another new
erhoben wurden. generic name (the 27th in the family and for which
he feared he might be called a splitter!) for his
Introduction and History plant the following year. The new name was
Slow Beginnings Cleistocactus, named for the feature that the
Little did Lemaire suspect in 1844 when he stamens almost close the flower.
described a “pretty Cactus recently blossomed in Although we now unequivocally accept the
the stove-house of Courant at Le Havre” that it genus it was not so for the first 40 years or so,
was one of the most widespread species of what even though other species with partially closed
would eventually become a very popular genus in flowers were discovered and described. In 1897,
cactus horticulture. He named it Cereus when Schumann published the first edition of his
baumannii after Mr Napoleon Baumann of Gesamtbeschreibung der Kakteen, he listed 5
Bolwilliers from whom he had received a plant. names that we now include in Cleistocactus
Napoleon and his brother Augustus owned a (baumannii, hyalacanthus, laniceps, parviflorus
nursery and they had recently imported the plant and smaragdiflorus) but continued to call them
from the Cordilleras. It seems their marketing Cereus. It was not until 1904 that Roland-
strategy was excellent as the plant rapidly spread Gosselin, in an obscure French journal,
to major collections within France so that transferred these species to Cleistocactus, based
Monville was able to observe the flowers soon on a manuscript left by the late Frédéric Weber.
thereafter. Fortunately, he was able to Perhaps because of the obscure nature of the
communicate this to Lemaire prior to the publication and its limited distribution the name
publication of volume 5 of L’Horticulteur still did not come into common usage and it was
148 Bradleya 34/2016

only with the publication of Volume 2 of The accepted without evaluating their potential as
Cactaceae by Britton & Rose in 1920 that the new synonyms. This, however, would have been a
name became more widely known, although they difficult task at the time since little or no
still only included the same names as had information was available on the distribution or
Schumann. variability of the species in habitat. That
information became more readily available in
The Numbers Game subsequent years and was able to influence the
It was not until the exploratory activities of treatment in The New Cactus Lexicon (Hunt et al.,
Backeberg, Cárdenas and Ritter in the mid 20th 2006). In that work only 24 taxa (17 species + 7
century that the genus began to grow. Cárdenas heterotypic subspecies) were fully recognised with
was primarily interested in expanding the cactus a further 5 species considered as provisional, all
flora of Bolivia thus he contributed 27 new names, placed in Cleistocactus.
all of which derive from single populations, but The most recent treatment of the family (Lodé,
failed to provide a systematic treatment of the 2015) claimed to be “The new classification of
genus. On the contrary, Backeberg’s forte was Cacti mainly based on molecular data ...”.
systematic treatment and, although he only However, the treatment of Cleistocactus lies
described 15 new taxa he did recognise that 4 somewhere between those of Anderson (2001) and
older names also belonged to the genus and made Hunt et al. (2006) in accepting 27 species. It
the appropriate combinations. In his final differs markedly from both, however, in accepting
treatment of the genus, published shortly before two of the species under Borzicactus Riccob. For
his death in 1966, he accepted 47 taxa in one of these (C. samaipatanus (Cárd.) Hunt) the
Cleistocactus and a further 7 in 3 genera now reason appears to be based on a misreading of
considered to be synonymous. Ritter was the last taxon names in the cladogram of Hernández-
of the three to publish a treatment, in 1980, and Hernández et al. (2011) since the authors did not
was extremely critical of Backeberg’s work include the species in their study.
considering his descriptions riddled with mistakes
and inaccuracies and his genera over-inflated. Limited Specific Characters
Even so he too proposed a large genus of 44 taxa The clear message that can be extracted from
along with 2 monotypic genera for taxa now this history is that, although it is relatively easy
included in Cleistocactus. Among those were 26 of to identify plants as belonging to the genus, it can
his own descriptions, only 10 of which were known be very difficult to define the specific limits, as
by name to Backeberg. In the background, perhaps is the case for all major genera of the
Buxbaum was giving serious consideration to the Cactaceae. In the treatment that follows I cannot
use of morphology in cactus taxonomy. Having claim to be very different since I have encountered
discovered what he perceived as significant this situation on several occasions when observing
differences in the shape of the flowers of C. the plants in habitat. Unfortunately, there are few
baumannii compared to those of the remainder of characters that can be used to separate the
the species he proposed to divide the genus into species since all have a similar growth form,
two subgenera and created the subgenus essentially the same flower shape (with some
Annemarnieria for those species differing from the obvious exceptions) and more-or-less identical
type (Buxbaum, 1956). fruits and seed. Those characters that I have
found useful include branching mode (basitonic
The Modern Era only vs. basi- and mesotonic), flower colour and
Since 2000, three major treatments of the flower inclination (erect, angled or horizontal).
Cactaceae have appeared (Anderson, 2001; Hunt These characters become especially useful when
et al., 2006; Lodé, 2015) each with a different encountering sympatric species. This is a
treatment of Cleistocactus. The first (Anderson, frustratingly frequent occurrence, particularly in
2001), and its subsequent edition in German the core of the range where it is common to find
(Eggli, 2005), was quite uncritical and accepted two species, and sometimes three, at a single
many of the species erected by Cárdenas, location.
Backeberg and Ritter with little or no re-
evaluation of their status. Although managing to Systematic Position and Circumscription
reduce the taxon count to 38, there were still Buxbaum (1958) placed Cleistocactus as a core
many names based on single populations. The member of his subtribe Borzicactinae within the
basis for this work appears to have been the tribe Trichocereae when creating a formal division
CITES Cactaceae Checklist (Hunt, 1999). of Schumann’s ‘Cereoideae’. He placed it alongside
Unfortunately 15 of the names listed in that work several other genera including Arequipa,
as provisionally accepted were treated as fully Bolivicereus, Borzicactus, Denmoza, Loxantho-
Bradleya 34/2016 149

cereus, Matucana, Morawetzia, Oreocereus, Oroya but, given the limited taxon sampling from within
and Seticereus based on their possession of Cleistocactus, the low number of markers
common floral traits. It is testament to Buxbaum’s employed and the lack of nuclear sequences I feel
careful and detailed analysis that this grouping it prudent to wait for further molecular data
has remained together throughout all subsequent before making further adjustments to the
modern classifications with no dispute. Although circumscription of the genus.
some of the genera have been absorbed, for
example Arequipa into Oreocereus, Cleistocactus Study Methods
has remained intact albeit at times taken to Names. An initial list of names was obtained
include species from the other genera. Perhaps by conducting a search of The International Plant
the earliest examples are the inclusion of Names Index (IPNI; 2012) for records from the
Echinocactus rhodacanthus by Lemaire in 1861 following genera: Cleistocactus, Bolivicereus,
and subsequently the inclusion of Oreocereus Hildewintera, Winteria, Winterocereus,
celsianus by Weber in 1904. More recently, Borzicactus and Loxanthocereus, and
following the deliberations of the IOS Working subsequently for the species epithets: strausii and
Party (1986), the scope of the genus was expanded baumannii. Over 300 records were identified as
to include all those cereoid species from Andean belonging to members of the genus Cleistocactus.
South America having ornithophilous flowers and These records were downloaded in extended
solid fruits. Thus many species of Borzicactus format and loaded into a searchable, indexed
(including Akersia, Borzicactella, Clistantho- spreadsheet (Libre Office Calc) for improved
cereus, Loxanthocereus, Maritimocereus and access. This file is available from the author on
Seticereus) subsequently gained combinations in request.
Cleistocactus through the publications of several Occurrence data. The primary dataset of
authors (Hunt & Taylor, 1987; Hunt, 2003; occurrence records was obtained several years
Ostolaza, 1996, 1998; Ostolaza et al. 2003; previously through a search of the Global
Leuenberger, 2002). This was the definition of Biodiversity Information Facility (www.gbif.org;
Cleistocactus used, with some reservation, in The last accessed 8th April 2009) for all records of the
New Cactus Lexicon (Hunt et al., 2006). family Cactaceae. This resulted in 21,538 records
At that time few studies using molecular which were incorporated into an SQL-based
techniques to investigate phylogeny had been database using custom written software. The
published and those available had not database was augmented with further records
concentrated on Cleistocactus and related genera. donated by many friends and travelling
It was not until 2012 that a study including a companions and is continually updated with
significant number of species from the occurrence information from my own field trips.
Borzicactinae was published (Schlumpberger & The number of records now stands at 36,768 of
Renner, 2012). The results clearly demonstrated which 420 relate to Cleistocactus.
that Cleistocactus as circumscribed by Hunt et al. Mapping. The 420 records of Cleistocactus
(2006) was polyphyletic. The genus appeared in a were exported from the database and converted
moderately well supported clade sister to a clade into ESRI shapefiles using the free software
containing only some members of Echinopsis s.l. DIVA-GIS (www.diva-gis.org). The package was
as well as Harrisia Britton, and distant from that used to generate point distributions for each
containing Borzicactus sensu Kimnach. The clade species using base maps built from country level
containing Cleistocactus s.s. was poorly resolved spatial data available at the same website. Shape-
and contained several other genera, specifically, files depicting the ecoregions of the world (Olsen
Cephalocleistocactus Ritter, Vatricania Backeb., et al, 2001) were obtained from the World Wildlife
Samaipaticereus Cárd., Weberbauerocereus Fund (www.worldwildlife.org/publica-tions/ter-
Backeb. and Yungasocereus Ritter. Thus the restrial-ecoregions-of-the-world; last accessed 9th
situation is very complex and currently only August 2016). These were overlaid on the base
partly resolved. A first step was taken by Hunt map and allowed identification of the major
(2013) in an update to The New Cactus Lexicon biome(s) and ecoregion(s) within which each
where Borzicatus was reinstated. A broader species can be found.
alternative view was taken by Anceschi & Magli
(2013) who, since the study also demonstrated Taxonomy
Echinopsis s.l. to be polyphyletic, transferred The following list describes all those taxa that
several species of Cleistocactus, including the I consider sufficiently distinct to require formal
type, into Echinopsis thus making the former a recognition. The descriptions are based on those
synonym. There are several other, less given in The New Cactus Lexicon (Hunt et al.,
consequential, options that could be considered 2006) with additional information included from

the protologues of the basionym and subsumed Etymology: From Greek kleistos, closed, and
synonyms where relevant. They are also expanded Cactus, referring to the way the throat of the
with information from my own observations, both flower is almost closed.
in the field and in culture, when details, for
example, of flowers and fruits, were not given in Habitat & Distribution: Arid and humid zones of
the first description. All references to the the chaco, inter-Andean valleys, Yungas and pre-
International Code of Nomenclature refer to the puna generally amongst low shrubs and bushes
Melbourne Code (McNeill et al., 2012). but occasionally in deciduous forest. Generally
terrestrial but frequently lithophytic or even
Cleistocactus Lem., Ill. Hort. 8: Misc. 35. (1861). epiphytic, 100–3700m.
Type: Cereus baumannii Lem. Ecological regions: The species are found in 10
Bolivicereus Cárd., Cact. Succ. J. (US) 23: 91 (May different ecoregions with major concentrations in
1951). Type: Bolivicereus samaipatanus Cárd. the Bolivian Montane Dry Forest and adjacent
Borzicactus subg. Bolivicereus (Cárd.) Buxbaum, Southern Andean Yungas. Other frequently
Kakteen (H. Krainz) 58: CVc (1974). Basionym: occupied regions include the Central Andean
Bolivicereus Cárd. Puna and Bolivian Yungas. One species occurs
Cephalocleistocactus Ritter, Succulenta (NL) extensively throughout the Dry & Humid Chaco
1959: 108 (1959). Type: Cephalocleistocactus whilst two others are found in the Peruvian
chrysocephalus Ritter. Yungas.
Hildewintera Ritter, Kakt. and. Sukk. 17: 11 Occurrence: AR (CC, CD, CT, FO, JY, LR, SA, SE,
(1966), nom. inval. [Art. 33.4]. Type: Winteria TM), BO (CB, CH, LP, PO, SC, TR), BR (MS), PE
aureispina Ritter. (AP, AY, CZ, HC, JU), PY (AG, BQ, CE, CN, CR,
Hildewintera Ritter ex G.D. Rowley, Regnum. GU, PG, PH), UY.
Veg. 54: 15 (1968), nom. illeg. [Art. 52.1]. Type:
Winteria aureispina Ritter. 1. Cleistocactus baumannii (Lem.) Lem., Ill.
Loxanthocereus subg. Hildewintera Buxbaum, Hort. 8: Misc. 35 (1861). Basionym: Cereus
Kakteen (H. Krainz) 58: CVb (1974). baumannii Lem.
Seticleistocactus Backeb., Descr. Cact. Nov. 3: 13 Aporocactus baumannii (Lem.) Lem. Ill. Hort. 7:
(1963). Type: Cleistocactus piraymirensis Misc. 66–68 (1860). Basionym: Cereus
Cárd. baumannii Lem.
Winteria Ritter, Kakt. and. Sukk. 13: 4–8, figs. Cereus anguinus Gürke, Monats. f. Kaktk. 17: 166
(1962), nom. illeg. [Art. 53.3]. Type: Winteria (1907). Type: Paraguay, (not preserved?).
aureispina Ritter. C. baumannii Lem., Hort. Univ. 5: 126, 315
Winterocereus Backeb., Kakteenlexikon p.455 (1844). Type: cult. hort. Paris from hort.
(1966). Type: Winteria aureispina Ritter. Baumann, not preserved. Neotype selected by
Hunt & Taylor, Cact. Syst. Init. 21: 6 (2006):
Shrubs, generally with several slender, Jardin Fleuriste 1: t.48 (1851).
cylindrical stems which can be erect, ascending, C. baumannii var. colubrinus K. Schum. (non C.
decumbent, procumbent or pendent. Stems with colubrinus Otto), Gesamtb. Kakt. p.133 (1898).
5–30 ribs and closely spaced areoles bearing few C. baumannii var. flavispinus K. Schum.,
to many acicular spines. Flowers produced along Gesamtb. Kakt. p.133 (1898).
the upper section of each stem below the apex C. colubrinus Otto, in Förster’s Handb. Cactkde.
normally from undifferentiated areoles but p.409 (1846).
occasionally along with extra bristles and spines. C. colubrinus var. flavispinus Salm-Dyck, Cact.
The flowers are tubular, regular to zygomorphic, Hort. Dyck. 1844: 32 (1845).
diurnal, and vary in colour from red to green. The C. grossei Weingart, nom. nud., Monats. f. Kaktk.
hypanthium, occasionally markedly kinked above 18: 8 (1908).
the ovary, is covered with many small scales. C. tweediei Hooker, Bot. Mag. 76: t.4498
Tepals scarcely larger than the scales and in most (1850).Cleistocactus anguinus (Gürke) Britt. &
species barely spreading, though if spreading then Rose, The Cactaceae 2 p.175 (1920). Basionym:
often obliquely. Stamens inserted in two series, Cereus anguinus Gürke.
included or exerted; anthers and pollen often C. aureispinus Frič, nom. nud., Möller’s Deutsche
purplish. Nectar chamber partially closed by Gärtn.-Zeitung xliii: 267 (1928).
lowermost filaments or by a diaphragm. Style C. baumannii ssp. anguinus (Gürke) Braun &
mostly exerted, generally white or yellow. The Esteves, Succulenta (NL) 74: 84 (1995).
fruits are small and globose retaining the floral Basionym: Cereus anguinus Gürke.
remains. They are solid with fleshy pulp enclosing C. baumannii ssp. chacoanus (Ritter) Braun &
small pyriform shiny black seeds. Esteves, Succulenta (NL) 74: 84 (1995).

C. chacoanus Ritter, Kakt. Südamer. 2: 671, Abb.
685 (1980). Type: Villa Montes, Prov. Gran
Chaco, Bolivia, Ritter 841 (U).
C. chacoanus var. santacruzensis (Backeb.) Ritter,
Kakt. Südam. 2: 672, Abb. 687 (1980).
Basionym: Cleistocactus santacruzensis
Backeb. Note: Ritter added further details
regarding the location, viz: “tiled roofs of house
in the capital”, Santa Cruz, Bolivia.
C. croceiflorus Ritter, Kakt. Südamer. 1: 272
(1979). Type: in the neighbourhood of Cerro
Gavan, Puerto Casado, Dept. Boquerón,
Paraguay, Ritter 1185 (U).
C. colubrinus var. flavispinus (Salm-Dyck) Borg,
Cacti p.150 (1937). Basionym: Cereus
colubrinus var. flavispinus Salm-Dyck.
C. colubrinus var. grossei Borg, nom. nud., Cacti
p.150 (1937).
C. flavispinus Backeb., Die Cactaceae II: 1000
(1959). Basionym: Cereus baumannii var.
flavispinus K. Schum.
C. grossei Backeb., nom. nud., Descr. Cact. Nov. I
p.17 (1957).
C. jugatiflorus Backeb., Die Cactaceae VI: 3685
(1962). Type: the cited illustration.
C. paraguariensis Ritter, Kakt. Südamer. 1: 273
(1979). Type: rocky hills near Paraguari, Dept
Figure 1. Cleistocactus baumannii LM0507.01 Paraguari, Paraguay, Ritter 1185b (U).
north of Eyti, Bolivia, 980m. The form previously C. santacruzensis Backeb., Kakteenlexikon 89, fig.,
known as ssp. santacruzensis. (1966). Type: Santa Cruz, Bolivia, Ritter 356
Photograph: Martin Lowry (U).
Echinopsis baumannii (Lem.) Anceschi & Magli,
Basionym: Cleistocactus chacoanus Ritter. Cactusinhabitat South America 2011/2013,
C. baumannii ssp. croceiflorus (Ritter) Braun & p.37 (2013). Basionym: Cereus baumannii
Esteves, Succulenta (NL) 74: 84 (1995). Lem.
Basionym: Cleistocactus croceiflorus Ritter. E. santacruzensis (Backeb.) Anceschi & Magli,
C. baumannii ssp. paraguariensis (Ritter) Braun Cactusinhabitat South America 2011/2013,
& Esteves, Succulenta (NL) 74: 84 (1995). p.40 (2013). Basionym: Cleistocactus
Basionym: Cleistocactus paraguariensis santacruzensis Backeb.
Ritter. Stems erect or arching to 2m × 1–4cm, branch-
C. baumannii ssp. santacruzensis (Backeb.) ing from the base with 12–17 ribs. Areoles ±1cm
Mottram, Cact. Succ. J. (US) 61: 156 (1989). apart bearing 15–20 acicular radial spines 1–2cm
Basionym: Cleistocactus santacruzensis long and 2–3 thicker central spines to 4cm. Flow-
Backeb. ers red-orange, markedly “S”-shaped, 5–7cm × 7–
C. baumannii var. colubrinus (Otto) Riccobono, 8mm, broadening to 1cm below the corolla.
Boll. Reale Orto Bot. Palermo 8: 266 (1909). Hypanthium with red scales each producing sev-
Basionym: Cereus colubrinus Otto. eral white hairs. Inner tepals red/orange, ex-
C. baumannii var. flavispinus (Salm-Dyck) panding slightly to reveal the filaments and style.
Riccobono, Boll. Reale Orto Bot. Palermo 8: The anthers produce red-magenta pollen. Fruit 1–
266 (1909). Basionym: C. colubrinus var. 1.5cm diameter, globose.
flavispinus Salm-Dyck.
C. baumannii var. grossei (Weingart) Marshall, Key to subspecies of C. baumannii
nom. nud., Cactaceae (Pasadena), p.106 1. Stems > 2cm diameter, spines > 1cm long
(1941). (AR, BO, S.PY) a. ssp. baumannii
C. bruneispinus Backeb., Die Cactaceae II: 1001 Stems < 2cm diameter, spines < 1cm long (BR,
(1959). Replaced synonym: Cereus baumannii N PY) b. ssp. horstii
var. colubrinus K. Schum.

Figure 2. Cleistocactus baumannii ssp. horstii at Fazenda Bala near Porto Murtinho, Brazil.
Photograph: Paulo Robson de Souza
a. ssp. baumannii Argentina. It is not completely clear from
Hooker’s text which of these was used for the
Stems erect, ascending or decumbent, > 2cm drawing, however, I consider it more likely to have
diameter. Spines 10–15, the centrals to 3cm long been the plants from the Hammersmith Nursery,
(Figure 1). which supplied Kew’s “largest and flowering
plant”, since the English description speaks of
Etymology: named after Mr Napoleon Baumann “Our tallest plants ...”.
of Bolwilliers (sic) who gave Lemaire the plant he The decision to include Backeberg’s C.
described. santacruzensis and synonyms here is taken after
observing several populations in habitat. The only
Habitat & Distribution: Shrubland along the character which appears to separate the two
Chaco margin amongst trees in loamy-sandy soils. forms is stem diameter, with those in the north
Its range is extensive reaching from La Rioja (Santa Cruz to Camiri), generally thinner than
Province, Argentina in the south to Santa Cruz de those in the south (La Rioja and Córdoba).
la Sierra, Bolivia in the north and extending Between these extremes (Villamontes to Santiago
eastwards from Villamontes in Bolivia, across del Estero) there are populations of intermediate
Paraguay to Asunción. 150–1050m. thickness.
Ecological regions: Dry Chaco, Chiquitano Dry
Forest and Humid Chaco. This is the only species b. ssp. horstii (P.J. Braun) N.P. Taylor, Cact.
adapted to the harsh arid conditions of these Cons. Init. 6: 15 (1998). Basionym: Cleistocactus
regions. horstii P.J. Braun.
Occurrence: AR (CC, CD, FO, LR, SA, SE), BO C. horstii P.J. Braun, Kakt. and. Sukk. 33: 204–
(CH, SC, TR), PY (AG, BQ, CE, CN, CR, GU, PG, 208, figs. (1982). Type: lowlands of Río
PH), UY. Map 1. Amoguija, SW of Mato Grosso do Sul, Brazil,
150m, 1974, Horst & Uebelmann 373
The plate chosen as the neotype for Cereus (KOELN).
baumannii Lem. is a copy of the illustration from Echinopsis horstii (P.J. Braun) Anceschi & Magli,
the protologue of Cereus tweediei Hooker. Kew Cactusinhabitat South America 2011/2013
received plants from the Messrs Lee of the p.38 (2013). Basionym: Cleistocactus horstii
Hammersmith Nursery and also grew plants from P.J. Braun
seeds from a Mr Tweedie of Buenos Aires, Stems decumbent to prostrate, to 1cm

Figure 3. Cleistocactus brookeae LM0512.02 east Figure 4. Cleistocactus brookeae LM0636.01 at
of Monteagudo, Bolivia, 1300m. Puente Azero, Bolivia, 1060m. The form previ-
Photograph: Martin Lowry. ously known as ssp. vulpis-cauda.
diameter, spines < 1cm long (Figure 2). Photograph: Martin Lowry.
Gesamtbeschreibung. From there we learn that
Etymology: named to honour Leopoldo Horst, Anisits collected the plant “bei Porongo”, an
Brazilian cactus explorer and friend of Pierre Estancia in Dept. Presidente Hayes, Paraguay,
Braun. some 240km south west of Porto Murtinho, the
type locality of C. horstii, and much nearer to
Habitat & Distribution: lowlands west of the other known locations of C. baumannii ssp.
Pantanal, 100m. baumannii. Secondly, at subspecies level, the
Ecological regions: Humid Chaco. epithet anguinus has 3 years priority over horstii.
Occurrence: BR (MS), PY (CN). Map 1. Making them synonymous would require a change
of name and introduce potential nomenclatural
Although the protologue of Cleistocactus confusion.
horstii suggests a sparsely-spined plant, there are
several clones of HU 373 in cultivation which 2. Cleistocactus brookeae Cárd, Cact. Succ. J.
demonstrate spination similar to that of the type (US) 24: 144, figs., (1952) (as ‘brookei’). Type:
of the subspecies. “Pozo no. 4”, Camiri, Prov. Cordillera, Dept. Santa
The remarkable similarity of Backeberg’s Cruz, Bolivia, 900m, Mar 1949, A. Corro s.n. in
illustration Abb. 921 in Die Cactaceae of what he Cárdenas 4818 (LIL 531590, holo.); between
believed to be Gürke’s Cereus anguinus with the Zudañez and Incahuasi, Prov. Hernando Siles,
photographs in the protologue of C. horstii P.J. Dept. Chuquisaca, 1200m, Aug 1949, Miss W.
Braun leads me to suspect that these might Brooke s.n. (US, para.).
represent the same taxon. I refrain from making C. brookeae var. flavispinus Ritter, Kakt.
them synonymous here for two reasons. First, Südamer. 2: 680, Abb. 664 (1980). Type: “some
Gürke provided no explicit type location with his km northwest from Boyuibe”, Prov. Cordillera,
description saying only that Schumann had listed Dept. Santa Cruz, Bolivia, 1963, Ritter 1129
it as Anisits 28 on page 41 of the Nachträge of his (U).

Etymology: named to honour Miss Winifred
Brooke who discovered the plant.
Habitat & Distribution: Steep cliffs and rocky

slopes close to rivers in the foothills of the eastern
Andes, 750–1300m.
Ecological regions: South Andean Yungas and
margins of Dry Chaco.
Occurrence: BO (CH, SC, TR). Map 2.
Although not rediscovered in habitat for many

years, since 2004 it has been observed at several
locations along the eastern margin of the Andean
foothills allowing a clearer assessment of its
variation. At the southern locations the plants are
generally erect, e.g. Ritter’s var. flavispinus,
whereas as one goes north they become
decumbent and eventually pendant. Hence the
former variety/subspecies vulpis-cauda is
included here since its main differentiating
character, pendant stems, is not unique and can
also be seen in several populations of the type.
The geniculate (bent like a knee) flower and
staminodial diaphragm in this species are
effective adaptations to ornithophily, both serving
to prevent insects from reaching the nectar
chamber. Similar adaptations are seen separately
in the two subspecies of C. winteri clearly linking
Figure 5. Cleistocactus brookeae GC857.02 from that species as a florally specialised member of
east of Monteagudo, Bolivia, in culture. Cleistocactus.
Photograph: Graham Charles.
3. Cleistocactus buchtienii Backeb., Kaktus
C. brookeae ssp. vulpis-cauda (Ritter & Cullmann) ABC pp.189, 411 (1936) Type: between Arque and
Mottram, Cact. Syst. Init. 13: 10 (2002). Cochabamba, Dept. Cochabamba, Bolivia, 2000m,
Basionym: C. vulpis-cauda Ritter & not preserved. Neotype (designated here):
Cullmann. Angostura, 2560m; Oct 1947, Cárdenas 4820a
C. vulpis-cauda Ritter & Cullmann, Kakt. and. (LIL 531560).
Sukk. 13: 38–40, figs. (1962). Type: Puente C. angosturensis Cárd., Cact. Succ. J. (US) 28: 60,
Acero, Prov. Tomina, Dept. Chuquisaca, figs. (1956). Type: Angostura, Mun. Tarata,
Bolivia, Ritter 847 (ZSS, not found). Lectotype Prov. Arce, Dept. Cochabamba, Bolivia,
(designated here): the specimen 124821 at 2550m, August 1955, Cárdenas 5055 (LIL
SGO labelled “Pte. Acero Nr. 12”. 531595).
C. wendlandiorum Backeb., Kakt. and. Sukk. 6: C. ayopayanus Cárd., Cact. Succ. J. (US) 28: 58,
113–117, figs. (1955). Type: fig.114, in Haage, figs. (1956). Type: between Morochata and
Freude mit Kakteen (1954), cited on p.117 of Independencia, Prov. Ayopaya, Dept.
the protologue. Cochabamba, Bolivia, 2300m, October 1949;
A shrub with stems erect or pendant, Cárdenas 5053 (LIL, not found; US, not
branching from the base to 2m × 2–4cm. Ribs 18– recorded). Lectotype (designated here): fig. 35
24 bearing areoles 3–4mm apart. Spines 25–40, of the protologue.
undifferentiated, about 1cm long, yellow or white. C. buchtienii var. flavispinus Cárd., Cact. Succ. J.
Flowers produced near stem apex, always vertical (US) 24: 182, fig. (1952). Type: La Angostura,
regardless of stem orientation, 5cm × 8mm, red or Mun. Tarata, Prov. Arce, Dept. Cochabamba,
orange. Hypanthium geniculate above ovary; Bolivia, 2570m, January 1948, Cárdenas
perianth opening obliquely, not expanded. Style 4820a (LIL 531560, holo; US 2342649, iso).
shortly exerted; stamens basally connate; anthers C. orthogonus Cárd., Cactus (Paris) xiv: 161
purple. Fruit globose, to 1cm diameter, purple (1959). Type: between Chaqui and Betanzos,
(Figures 3–5). Prov. Saavedra, Dept. Potosi, Bolivia, 2800m,
January 1956, A. Vidaurre in Cárdenas 5500

Figure 6. Cleistocactus buchtienii LM0452.02 Figure 7. Cleistocactus buchtienii ‘tupizensis’
south of Kami, Bolivia, 3300m. The form previ- GC892.01, Road to Culpina, Bolivia 3120m.
ously known as C. ayopayanus. Photograph: Graham Charles.
Photograph: Martin Lowry.
and higher, 5cm diameter. Ribs 12–18, low,
(LIL 531581, US). Note: the altitude stated in slightly grooved above the areole. Areoles < 1cm
the protologue is not possible at the stated apart, tomentose, with 10–15 spines, barely
location and must be an error since the differentiable, acicular to 2.5cm long. Flowers
holotype label at LIL reads “3400m” which is sparingly from upper part of stem, tubular, 5–8cm
correct. × 9mm, directed upwards at ~45° to the stem,
C. ressinianus Cárd., Cact. Succ. J. (US) 28: 55, deep red–purple with exerted green stigma and
figs., (1956). Type: between Sucre and purple anthers. Fruit globular, ±1cm diameter,
Tarabuco, Prov. Oropeza, Dept. Chuquisaca, red. Seeds, 1.2mm, black, pitted (Figure 6).
Bolivia, 2700m, February 1949, Cárdenas
5051 (LIL 531567). Etymology: named to honour Otto Buchtien who
C. sucrensis Cárd., Cact. Succ. J. (US) 24: 148. collected cacti in Bolivia in 1932 and 1934.
(1952). Type: Takos, “kilometre 7 from Sucre
to Tarabuco”, Prov. Oropeza, Dept. Habitat & Distribution: Sparsely wooded slopes
Chuquisaca, Bolivia, 2750m, February 1949, and plains in arid inter-Andean valleys in the
Cárdenas 4820 (LIL 531572, US). North. The southern form generally in more arid
C. tupizensis sensu Backeb., Kakteen-Freund 3: locations, 2200–3700m.
124 (1934), non Cereus tupizensis Vaupel. Ecological regions: Bolivian Montane Dry Forest
C. tupizensis var. sucrensis (Cárd.) Backeb., Die and Central Andean Puna.
Cactaceae II: 1012 (1959). Basionym: Occurrence: BO (CB, CH, PO, TR). Map 3.
Cleistocactus sucrensis Cárd.
Echinopsis buchtienii (Backeb.) Anceschi & Magli, The name as now used includes not only
Cactusinhabitat South America 2011/2013 plants from the region of the type locality but also
p.37 (2013). Basionym: Cleistocactus those from populations from much further south
buchtienii Backeb. for which the misapplied name C. tupizensis
Stems erect to 2m, branching from the base (Vaupel) Backeb. was once used. Since the

original observation by Mottram (2002), the
basionym of this combination has now been
conclusively demonstrated (Leuenberger, 2012) as
a synonym of Oreocereus celsianus (Salm-Dyck)
Riccobono. The plants to which the name C.
tupizensis have been misapplied show
considerable differences from the type. They are
generally shorter in stature, have more ribs and
more, often hyaline spines giving them a silver
appearance. The flowers are also much shorter (4–
5cm vs 8–9cm) with fewer hairs on the
hypanthium (Figure 7). These characters give the
plants an appearance similar to that of C.
hyalacanthus (K. Schum.) Rol.-Goss. and it may
be better to consider these southern populations
under that name rather than here.
Although Backeberg declared no type
specimen, the New Cactus Lexicon (NCL) records
Eggli as documenting that a specimen at ZSS
collected by Buchtien can serve as the type. I can
find no record of this specimen in the herbarium
database of ZSS hence, given the above
discussion, it seems desirable to provide an
unambiguous typification. Backeberg provided no
illustrations in the protologue that could serve as
a lectotype and subsequent illustrations, for
example, in Die Cactaceae, are insufficiently clear.
Of the subsequent names now considered
synonymous, only two are from within a Figure 8. Cleistocactus candelilla BCMdV206.03
reasonable proximity of Backeberg’s quoted type from south of Mataral, Bolivia, 1670m in culture.
locality. These are C. angosturensis and C. Photograph: Martin Lowry.
buchtienii var. flaviflorus, both names published
by Cárdenas with type sheets present at LIL. designated. Lectotype (designated here): Fig.
Since the latter was obviously considered closer to 95 of the protologue.
C. buchtienii Backeb. by Cárdenas, its specimen C. muyurinensis Ritter, Taxon 13: 114 (1964).
is more representative and there is an excellent Type: Muyurina, Prov. Valle Grande, Dept.
photograph in the protologue so it is an Santa Cruz, Bolivia, Ritter 821 (U 117562B).
appropriate choice as neotype. Although the type C. piraymirensis Cárd., Cact. Succ. J. (US) 33: 78,
sheet does not contain flowers, a description in figs. (1961). Type: Río Piraymiri, Prov. Valle
Cárdenas’s handwriting states “wine red, long” on Grande, Dept. Santa Cruz, Bolivia, 1800m,
the label and the photograph shows them rising Sep 1959, Cárdenas 5526 (LIL, not found).
at an acute angle from the stem, all significant Lectotype (designated here): Fig. 40 of the
characters in identifying this species. protologue.
C. pojoensis (Cárd.) Backeb., Die Cactaceae II:
4. Cleistocactus candelilla Cárd., Cact. Succ. 1003 (1959). Basionym: C. candelilla var.
J. (US) 24: 146–147, figs., (1952). Type: La Tigre pojoensis Cárd.
(Pampa Grande), Prov. Florida, Dept. Santa Cruz, Echinopsis candelilla (Cárd.) Anceschi & Magli,
Bolivia, 1400m, Oct 1950, A. Corro s.n. in Cactusinhabitat South America 2011/2013
Cárdenas 4819 (LIL, not found; US, not recorded). p.38 (2013). Basionym: Cleistocactus
Lectotype (designated here): Fig. 94 of the candelilla Cárd.
protologue. Seticleistocactus piraymirensis (Cárd.) Backeb.,
C. candelilla subsp. piraymirensis (Cárd.) Descr. Cact. Nov. 3: 13 (1963). Basionym: C.
Mottram, Cact. Syst. Init. 13: 10 (2002). candelilla var. piraymirensis Cárd.
Basionym: C. piraymirensis Cárd. C. vallegrandensis Cárd., Cact. Succ. J. (US) 33:
C. candelilla var. pojoensis Cárd., Cact. Succ. J. 75–76, figs. (1961). Type: between Vallegrande
(US) 24: 147 (1952). Type: Hacienda La and Candelaria, Prov. Valle Grande, Dept.
Habana (Pojo), Prov. Carrasco, Dept. Santa Cruz, Bolivia, 2600m, October 1959;
Cochabamba, Bolivia, 2500m; specimen not Cárdenas 5523 (LIL, not found). Lectotype

Figure 9. Cleistocactus candelilla LM0730.02
east of Mataral, Bolivia, 1365m.
Figure 12. Cleistocactus capadalensis LM0768.01

west of the Rio Pilcomayo near Kollpa, Bolivia,
2120m. Photograph: Martin Lowry.
Figure 10. Cleistocactus candelilla LM0736.01

east of Pena Colorado, Bolivia, 1675m. A form
with exceptionally hairy flower tubes.
Figure 11. Cleistocactus capadalensis LM0865.01 Figure 13. Cuttings of Cleistocactus chryso-
east of Capacdala, Bolivia, 2015m, near the type cephalus from near Inquisivi, Bolivia.
locality. Photograph: Martin Lowry. Photograph: Holger Kelsch

(designated here): fig. 34, of the protologue.
Syn. nov. Originally described without details of flower
A small shrub to 1m tall, branching from the and fruit this taxon was long considered a
base. Stems, generally many, erect, ascending or synonym of C. buchtienii Backeb. until
decumbent, 2–3cm diameter with 10–15 ribs rediscovered in 2012 close to the type locality by
bearing areoles < 1cm apart. Spines the author and Mats Winberg. The description
differentiated; radials, 12–18, yellow to white, up above is expanded from that of Ritter to include
to 1cm long; centrals, 1–4, yellowish brown, 10– details of the flowers and fruit observed in
25mm. Flowers, straight, 3–6cm × 4–7mm habitat. It was not recognized in the NCL and
diameter; hypanthium magenta, sometimes consequently its conservation status was not
covered with white hairs to 1cm; outer tepals assessed. Given its small area of extent and area
somewhat expanded, magenta or yellowish, inner of occupancy with the threats due to low level
tepals magenta. Fruit globose, orange-red to subsistence farming it is probably best to consider
purple, ±12 mm (Figures 8–10). it as Vulnerable.
Etymology: the Spanish vernacular name for the 6. Cleistocactus chrysocephalus (Ritter)
plant meaning “small candle” and apparently Mottram, Cact. Syst. Init. 13: 11 (2002).
referring to the flame-like flowers. Cephalocleistocactus chrysocephalus Ritter,
Succulenta (NL) 1959: 107–111 (1959). Type:
Habitat & Distribution: dry semi-deciduous thorn gorge below Inquisivi, Prov. Inquisivi, Dept.
scrub in the higher interior valleys, 1300–2800m. La Paz, Bolivia, 2000m, October 1954, Ritter
Ecological regions: Bolivian Montane Dry Forest. 326 (ZSS).
Occurrence: BO (CB, CH, SC). Map 3. A tall shrub of 2–5m branching both from the
base and higher. Stems 3–5cm diameter with 11–
A very frequent plant in the dry thorn-scrub of 14 ribs bearing areoles 5–8mm apart. Spines
the north-eastern Andes. In the west its range differentiated; radials about 15, pale, about 7mm
overlaps with that of C. parviflorus (K. Schum.) long; centrals to 6, yellowish, 7–30mm long.
Rol.-Goss. where in many places they grow Flowers produced from a lateral pseudocephalium
sympatrically. It is possible that some inter- up to 1m long and 3–4cm wide densely packed
gradation of the two species occurs giving rise to with spines and bristles to 3cm long. The ribs in
plants with mixed characteristics (Figure 28). this zone reduced and the areoles closer together.
Flowers ±straight, 5cm long, yellow to red. Fruit
5. Cleistocactus capadalensis Ritter, Kakt. greenish-red, 2cm long × 3cm broad. Seeds small,
Südam. 2: 677 (1980). Type: slopes above Río 0.7mm, black, shiny, nearly smooth (Front cover
Pilcomayo, Capadala, Prov. Azurduy, Dept. picture and Figures 13–15).
Chuquisaca, 1958, Ritter 828 (U).
Etymology: from Greek; “golden-headed”,
Stems ascending or decumbent 1–1.5m long × referring to the colour of the cephalium.
3–5cm diameter branching from the base. Ribs,
13–18, bearing areoles 5–8mm apart. Spines 8– Habitat & Distribution: Sparsely wooded slopes of
12, bright yellow, the centrals, 2–3, thicker and seasonally dry valleys of the Yungas, 1900–
longer, to 3cm. Flowers straight, directed upwards 2500m.
at a slight angle, 2.5–3.5cm × 8mm. Receptacle Ecological regions: Bolivian Yungas.
green; hypanthium yellow-orange; inner petals Occurrence: BO (CB, LP). Map 9.
red. Stigma-lobes green, slightly exerted; anthers
magenta. Fruit globose, approx. 1cm diameter Originally considered by Ritter as belonging to
with persistent floral remains, initially green a separate genus, Cephalocleistocactus, this
ripening to red. Seeds unknown (Figures 11 & 12). unusual plant has rarely been seen in habitat.
There has however, been one recent report
Etymology: referring to the place at which Ritter (Kelsch, 2016) that it still exists near the type
discovered the species: Capadala in the gorge of locality. Ritter’s criteria for separating the genus
the Río Pilcomayo. were based on a longer and wider nectar chamber
than found in Cleistocactus and on the cephalium.
Habitat & Distribution: Sparsely wooded slopes Although exceptional, the cephalium as described
on both sides of the Río Pilcomayo in sandy soil, by Ritter is not a true cephalium as in the sunken
2000–2500m. cephalium of Espostoa Britton & Rose and should
Ecological regions: Bolivian Montane Dry Forest. still be considered a pseudo-cephalium. Indeed, it
Occurrence: BO (CH). Map 4. is possible that it represents a closer link to

Figure 14. Cleistocactus chrysocephalus from near Inquisivi, Bolivia, 2430m.
Photograph: Holger Kelsch
Figure 16. Cleistocactus dependens LM0668.01

Figure 15. Cleistocactus chrsocephalus Ritter 326 east of Samaipata, Bolivia, 850m.
in culture. Photograph: Graham Charles. Photograph: Martin Lowry

green. Filaments white, anthers purple. Style
greenish-white, slightly exerted. Fruit globular,
approx. 1cm diameter, deep red. Seeds 1mm long,
shiny, black, finely pitted (Figures 16 & 17).
Etymology: from the Latin dependens; referring to

the hanging nature of the stems.
Habitat & Distribution: Hanging from cliffs in the

gorges of the north-easternmost rivers of the
Andes, 800–1500m.
Ecological regions: Bolivian Montane Dry Forest.
Occurrence: BO (SC). Map 5.
Although the NCL originally considered it a

disjunct form of C. candelilla Cárd., it can easily
be distinguished by its much thicker, decumbent
to sprawling branches, rounded ribs, fewer,
shorter and thicker spines and flowers with green
inner tepals. It has recently (2011) been recorded
in the region of the type locality by the author,
John Carr and Moises Mendoza.
Backeberg transferred it to his genus
Seticeistocactus based on the presence of the
conspicuous hairs on the floral tube. However,
this feature is found sporadically throughout the
Figure 17. Cleistocactus dependens HNT57784 in genus and cannot be used as a generic character.
culture showing the hairs and bristles on the hy-
pantheum. Photograph: Graham Charles. 8. Cleistocactus hildegardiae Ritter, Kakt.
Südam. 2: 685, figs., (1980). Type: Caña Cruz,
Vatricania Backeb. since recent molecular Prov. Mendez, Dept. Tarija, Bolivia, 1962, Ritter
evidence (Schlumpberger & Renner, 2012) has 1126 (U).
placed this monotypic genus closer to Borzicactus janae Halda & Horacek, Cactaceae
Cleistocactus than to Espostoa. Similar, though etc. (Bratislava) 13: 17–19 (2003). Type:
less extreme, pseudocephalia are produced by a Estancia Caña Cruz, Prov. Mendez, Dept.
small number of other species in the genus (C. Tarija, Bolivia, 3200 m, December 2002;
ritteri, C. variispinus and C. viridiflorus). All of Halda & Horacek s.n. (PRC 4723). Syn. nov.
them are found in similar habitats in the Yungas A small shrub with erect stems freely
on the northern edge of the Andes (Map 9). branching from the base. Stems to 20–40cm × 1–
2cm diameter with 15–19 ribs bearing close-set
7. Cleistocactus dependens Cárd., Cact. Succ. areoles. Spines 18–28, acicular, yellow to red-
J. (US) 24: 143–144, figs. (1952). Type: “humid brown, radials numerous, 3–8mm; centrals 5–8,
rocky slopes”, between La Angostura and La thicker than radials, 1–3cm long. Flowers tubular,
Negra, Prov. Florida, Dept. Santa Cruz, Bolivia, straight, 2.5–3.5cm × 5–6mm; magenta to purple,
1200m, Oct 1950, A. Corro in Cárdenas 4817 (LIL, outer tepals spreading. Fruit globose, 1cm
not found; US 2906781, iso; UC, paraiso). diameter (Figure 18).
Seticleistocactus dependens (Cárd.) Backeb.
Kakteenlexicon p.406 (1966). Basionym: Etymology: named to honour Frau Hildegard
Cleistocactus dependens Cárd. Winter, Ritter’s sister who distributed his
Stems dependent, procumbent or clambering collected seeds.
to 2m, 3–4cm diameter, branching basally, dull
green. Ribs 10–12, rounded bearing areoles 1– Habitat & Distribution: dry high altitude valleys
1.5cm apart. Radial spines 8–12, 2–4mm, acicular, sides in stony soils, 2600–3300m.
grey. Central spines 1–3, 1–1.5cm, thicker and Ecological regions: Bolivian Montane Dry Forest,
darker. Flowers 4–4.5cm × 0.7cm, straight; Central Andean Puna and margins of Southern
pericarpel dark purple, hypanthium strawberry Andean Yungas.
red, both with scales bearing conspicuous short Occurrence: BO (CH, TR). Map 5.
white hairs and bristles; distal perianth-segments

Figure 18. Cleistocactus hildagardiae LM0546.02 Figure 19. Cleistocactus hyalacanthus
in the Paicho valley north of Iscayachi, Bolivia, LM0602.01 north of Jucanas, Bolivia, 2260m
3170m. Photograph: Martin Lowry. Photograph: Martin Lowry.
Figure 20. Cleistocactus hyalacanthus GC159.01, Figure 21. Cleistocactus laniceps LM0458.02
the form previously known as C. jujuyensis, from south-east of Independencia, Bolivia, 2110m
El Alisal, Quebrada del Toro, Salta, Argentina in showing the woolly buds after which it is named.
culture. Photograph: Graham Charles. Photograph: Martin Lowry.

This is the smallest member of the genus and Habitat & Distribution: Sparsely wooded hills and
at one time was regarded a synonym of C. cliffs of the eastern Andean mountains, 1700–
hyalacanthus (K. Schum.) Rol.-Goss. but now 3100m.
considered distinct since the two species have Ecological regions: Bolivian Montane Dry Forest,
been found growing sympatrically at several Central Andean Puna, High Monte and margins
locations. Originally known only from its type of Southern Andean Yungas.
locality, the Río Paicho valley in Tarija, but Occurrence: AR (JY, SA), BO (CH, TR). Map 8.
recently found by the author and Mats Winberg
on the north side of the Río Pilaya in Chuquisaca. One of the oldest names in the genus but
recently much confused by application to the
9. Cleistocactus hyalacanthus (K. Schum.) plants discussed under C. buchtienii Backeb. as
Rol.-Goss., Bull. Soc. Centr. Agric. Hort. Acclim. the southern form. These two plants can look very
Nice 44: 33 (1904). similar when not in flower. C. hyalacanthus
Cereus hyalacanthus K. Schum., Gesamtb. Kakt. generally produces many flowers simultaneously
p.101 (1897). Type: Prov. Jujuy, Argentina, whilst on C. buchtienii there are normally only a
Kuntze s.n., not preserved. few produced together. The flowers of the latter
Cleistocactus hyalacanthus ssp. tarijensis (Cárd.) are also more straight, longer (> 4cm) and purple
Mottram, Cact. Syst. Init. 13: 11 (2002). rather than red. When not in flower
Basionym: C. tarijensis Cárd. distinguishing them is difficult, however spination
C. jujuyensis (Backeb.) Backeb., Beitr. can help as plants of the southern C. buchtienii
Sukkulentenk. Sukkulentenpflege 1942: 49–50 tend to have centrals and radials of different but
(1942). Basionym: C. strausii var. jujuyensis more or less constant length whilst those of C.
Backeb. hyalacanthus are quite variable. Otherwise
C. jujuyensis var. fulvus Ritter, Kakt. Südamer. 2: location may be the only clue; plants of C.
480 (1980). Type: Purmamarca, Prov. Jujuy, hyalacanthus are plants of the moister eastern
Argentina, 1959, Ritter 59b (U). slopes whereas C. buchtienii prefers the drier
C. strausii var. fricii (Därfl.) Backeb., Die valleys further west.
Cactaceae II: 1015 (1959). Basionym:
Pilocereus strausii var. fricii Därfl. 10. Cleistocactus laniceps (K. Schum.) Rol.-
C. strausii var. jujuyensis Backeb., Cact. J. Goss., Bull. Soc. Centr. Agric. Hort. Acclim. Nice
(London) 3: 25 (1934). Type: Jujuy, Argentina. 44: 32 (1904).
C. tarijensis Cárd., Cact. Succ. J. (US) 28: 54 Borzicactus laniceps (K. Schum.) Backeb., Cact.
(1956). Type: low hills near Tarija city, Succ. J. (US) 3: 132 (1932). Basionym: Cereus
Cercado, Dept. Tarija, Bolivia, 1900m, Mar laniceps K. Schum.
1952; Cárdenas 5050 (LIL 531575, holo., US, Cereus laniceps K. Schum., Gesamtb. Kakt. p.93
iso.). (1897). Type: near Tunari, Dept. Cochabamba,
Demnosa straussii var. luteispina Frič, Möller’s Bolivia, 1300m, Kuntze (B, not preserved?).
Deutsch. Gärtn.-Zeit. 44: 170 (1929). Type: C. laniceps var. plurispinus Ritter, Kakt.
Argentina. Südamer. 2: 673 (1980). Type: below Inquisivi,
Echinopsis nothohyalacantha Anceschi & Magli, Prov. Inquisivi, Dept. La Paz, Bolivia, 1957,
Cactusinhabitat South America 2011/2013 Ritter 603a (U).
p.38 (2013). Replaced synonym: Cereus A shrub to 4m with erect stems branching
hyalacanthus K. Schum. from the base or higher. Stems green, 3–5cm
Pilocereus strausii var. fricii Därfl., Monats. diameter with ±9 ribs about 1cm high with cross-
Deutsch. Kakteen-Ges. 2: 55, fig. (1930). furrows, the areoles large, 6mm, and well
A low shrub with erect stems to 1m tall separated. Radial spines few, short; centrals,
branching from the base. Stems 4–6cm diameter when present, to 2cm, yellow when new fading to
with 15–20 ribs bearing areoles 5–8mm apart. grey. Flowers from lateral areoles, funnelform,
Spines 20–30, unequal, up to 3cm long, white, 3.5cm, red, receptacle with copious brown wool.
yellow or brown. Flowers in profusion from near Fruit globose, ±1cm diameter (Figure 21).
stem apex, directed slightly upwards, 3.5–4cm ×
5–7mm, slightly curved, hypanthium red, tepals Etymology: from the Latin for “woolly-head” and
slightly spreading, magenta. Fruit globose, about referring to the hairy covering of the receptacle.
12mm diameter, red to orange. Seeds black, shiny
(Figures 19 & 20). Habitat & Distribution: Dry valleys of the
northern Andes below the Yungas forests, 2100–
Etymology: from the Latin hyalinus and acanthus 2400m.
meaning “glassy-spined”. Ecological regions: Bolivian Montane Dry Forest.

Aramani, Prov. Murillo, Dept. La Paz, Bolivia,
October 1954.
A shrub to 3m with numerous erect stems,
branching primarily from the base. Stems 4–6cm
diameter with about 19 ribs bearing areoles ±1cm
apart. Spines 16–22, white, yellow or pale brown,
5–15mm long, the centrals, 1 or 2, the longest.
Flowers tubular, ±straight, 3cm × 5mm, red. Fruit
globose to 2cm diameter, salmon pink (Figure 22).
Etymology: referring to the place of its first

discovery, Luribay, a small town in southern
Department La Paz.
Habitat & Distribution: Northern dry inter

Andean valleys west of the Yungas, 2500–3600m.
Ecological regions: Bolivian Montane Dry Forest.
Occurrence: BO (LP). Map 6.
Included here are three taxa described by

Ritter which have not so far been rediscovered in
habitat and for which no material is known in
cultivation. The only illustrations Ritter
published for C. glaucus (Kakt. Südamer. 2: Abb.
667 & 668) do suggest a close relationship with C.
luribayensis. Unfortunately, Ritter himself never
published an illustration for C. granjaensis; he
does, however, refer to a photograph by Backeberg
Figure 22. Cleistocactus luribayensis LM0447.02 (Die Cactaceae VI: 3689 Abb. 3350b) which he
at Luribay, Bolivia, 2580m. concedes may be the correct plant observing
Photograph: Martin Lowry. though that it does not branch as high as
illustrated when seen in habitat. Backeberg also
Occurrence: BO (CB, LP). Map 6. published side-by-side black and white
photographs (Die Cactaceae II: 1022 Abb. 954 &
A very distintive species because of its very 955) of both of Ritter’s plants under their
few spines and funnelform flower. These giving collection numbers, which reveal significant
rise to the, here discounted, suggestion that it is a differences between them. He considered C.
natural hybrid between C. luribayensis Cárd. and granjaensis close to C. brookeae Cárd. Indeed, if
Yungasocereus inquisivensis Ritter. the plant illustrated in the colour photograph
(Abb. 3350b) really is Ritter 106 then it bears little
11. Cleistocactus luribayensis Cárd., Cact. resemblance to plants of C. luribayensis seen by
Succ. J. (US) 28: 59–60, figs., (1956). Type: the author at the type locality and would be
Luribay slopes, Prov. Loayza, Dept. La Paz, worthy of separate recognition.
Bolivia, 2600m, Apr 1955, Cárdenas 5054 (LIL,
not found; US, not found). Lectotype (designated 12. Cleistocactus micropetalus Ritter, Kakt.
here): fig. 37 of the protologue. Südamer. 2: 675, figs., (1980). Type: Concepción,
C. glaucus Ritter, Taxon 13: 114 (1964). Type: Prov. Avilez, Dept. Tarija, Bolivia, 1958, Ritter
Tirata, Prov. Murillo, Dept. La Paz, Bolivia, 830, (U).
Ritter 112 (U 117611B). C. clavicaulis Cárd., Cactus (Paris) No. 80-81: 20
C. glaucus var. plurispinus Ritter, Taxon 13: 114 figs (1964). Type: near Angosto de Villa
(1964). Type: border of provinces Loayza and Montes, Prov. Gran Chaco, Dept. Tarija,
Murillo, Dept. La Paz, Bolivia, Ritter 112a (U Bolivia, 460m, May 1958, Cárdenas 5525 (LIL,
117754B). not found). Lectotype (designated here): the
C. granjaensis Ritter, Kakt. Südamer. 2: 687 photograph with the protologue.
(1980). Type: La Granja, Prov. Murillo, Dept. C. crassicaulis Cárd., Cact. Succ. J. (US) 33: 77,
La Paz, Bolivia, 3000m, 1953, Ritter 106 (U, figs., (1961). Type: near Angosto de Villa
not found). Lectotype (designated here): the Montes, Prov. O’Connor, Dept. Tarija, Bolivia,
specimen T4477 at ZSS collected by Ritter at 460m, March 1952, Cárdenas 6124 (LIL, not

Figure 23. Cleistocactus micropetalus LM0520.02 Figure 24. Cleistocactus micropetalus GC863.02
west of Palos Blancos, Bolivia, 1020m. The form south of Palos Blancos, Bolivia, 880m.
previously known as C. crassicaulis. Photograph: Graham Charles.
found). Lectotype (designated here): fig. 38 of Etymology: from Latin referring to the tiny tepals.
the protologue.
C. crassicaulis var. paucispinus Ritter, Kakt. Habitat & Distribution: Lowlands and inter
Südamer. 2: 674–675 (1980). Type: bottom of Andean valleys of southern Bolivia, west of the
gorge N of Cajas, Prov. Mendez, Dept. Tarija, Chaco margin, 750–2300 m.
Bolivia, Ritter 829a (U). Ecological regions: Southern Andean Yungas, Dry
C. tominensis ssp. micropetalus (Ritter) Mottram, Chaco and Central Andean Puna.
Cact. Syst. Init. 13: 12 (2002). Basionym: Occurrence: BO (TR). Map 3.
Cleistocactus micropetalus Ritter.
C. viridialabastri Cárd., Cact. Succ. J. (US) 35: The NCL treatment considers it a subspecies
201, figs. (1963). Type: between Villa Montes of C. tominensis (Weingart) Backeb. However, I
and Palos Blancos, Prov. Gran Chaco, Dept. consider it sufficiently different through its fewer
Tarija, Bolivia, 600m, May 1959; Cárdenas ribs and fewer, less conspicuous radial spines that
5549 (LIL, not found; US, not recorded). allow sight of the body justifying its recognition
Lectotype (designated here): the photograph in as a distinct species. It has a well-defined
the protologue. distribution in the Andean foothills of central
An erect shrub branching from the base with Tarija at altitudes generally less than those of C.
stems 1.5–3m tall. Stems 6–8cm diameter with tominensis. Further, C. tominensis is less clearly
16–18 ribs bearing areoles ±1cm apart. Spines defined, particularly at its northern border, and it
acicular, golden yellow; radials 5 or 6, 5–15mm is therefore better to maintain C. micropetalus as
long; centrals at least 1, 2–4cm. Flowers straight, distinct to avoid confusion.
tubular, 2–3cm, perianth rarely expanding;
hypanthium green becoming pinkish at anthesis,
covered by short white hairs. Fruit globose, 15–
20mm diameter, yellowish (Figures 23 & 24).

Figure 25. Cleistocactus morawetzianus Figure 26. Cleistocactus morawetzianus in cul-
LM0354.01 at Villa Azul, Peru, 2590m. ture showing the characteristic downward curva-
Photograph: Martin Lowry. ture of the flower. Photograph: Martin Lowry.
Figure 28. Cleistocactus aff. parviflorus

LM0839.01 east of Pasorapa, Bolivia, 1370m. An
Figure 27. Cleistocactus parviflorus LM0584.03 especially hairy-flowered form that may be a hy-
north of Vallegrande, Bolivia, 2000m. brid with C. candelilla.
Photograph: Martin Lowry. Photograph: Martin Lowry.

13. Cleistocactus morawetzianus Backeb., 14. Cleistocactus parviflorus (K. Schum.) Rol.-
Jahrb. Deutsch. Kakteen-Ges. 1: 77, fig. (1936). Goss., Bull. Soc. Centr. Agric. Hort. Acclim. Nice
Type: near Mariscal Cáceres, Ayacucho, Peru (fide 44: 32 (1904).
Backeb. Die Cactaceae 2: 1008 (1959)) (ZSS?). Cereus areolatus K. Schum., Gesamtb. Kakt. p.100
C. morawetzianus var. pycnacanthus Rauh & (1897). Type: mid-Andean region of South
Backeb., Descr. Cact. Nov. I: 17 (1957). Type: America, Kuntze s.n. (B†?).
near Ayacucho, Peru, 2700m, 1954, Rauh K75 C. parviflorus K. Schum., Gesamtb. Kakt. pp.100–
(ZSS). 101 (1897). Type: near Parotani, SW of
C. pycnacanthus (Rauh & Backeb.) Backeb., Cochabamba, Bolivia, 2400m, Kuntze s.n.
Kakteenlexikon p.88 (1966). Basionym: C. (B†?).
morawetzianus var. pycnacanthus Rauh & Cleistocactus areolatus (K. Schum.) Riccob., Boll.
Backeb. Ort. Bot. Palermo viii: 264–265 (1909).
C. villaazuensis Ritter, Kakt. Südamer. 4: 1359 Basionym: Cereus areolatus K. Schum.
1360, fig. (1981). Type: Villa Azul, Dept. C. fusiflorus Cárd., Cactus (Paris) No. 57: 252–
Huancavelica, Peru, 2000m, September 1964, 254, figs (1957). Type: near Saipena, Prov.
Ritter 1296, (U). Valle Grande (sic, actually Prov. Caballero),
A shrub to 2m with erect stems branching Dept. Santa Cruz, Bolivia, 1900m, April 1957,
from the base. Stems grey-green, ~5cm diameter Cárdenas 5491 (LIL, not found; US, not
with 12–14 ribs bearing areoles ±1cm apart, recorded). Lectotype (designated here): the
distinctly cross-furrowed. Spines 10–14, unequal, photograph in the protologue.
the longest to 1.5–5cm, yellow at first becoming C. herzogianus Backeb., Blätt. Kakteenf. Pt. 6, p.4
grey. Flowers 6–8cm × 9mm, white or green (1934). Type: “in Quebradas”, Prov. Arque,
occasionally becoming tinged with pink at Dept. Cochabamba, Bolivia, 2500m.
anthesis, distinctly down-curved, style exerted; C. ianthinus Cärd., Cact. Succ. J. (US) 28: 57–58,
self-fertile. Fruit globose with small scales, ~1cm figs (1956). Type: “on the Río Caine mountain
diameter, brownish-green. Seeds 1.2 × 0.8mm, slopes”, near Calahuta, Prov. Charcas, Dept.
keeled, shiny black (Figures 25 & 26). Potosi, Bolivia, 2000m, November 1949,
Cärdenas 5086 (LIL, not found; US, not
Etymology: named to honour Victor Morawetz, an recorded). Lectotype (designated here): fig. 33,
American philanthropic lawyer who financed two of the protologue.
of Backeberg’s expeditions to South America. C. parviflorus var. aiquilensis Ritter, Kakt. and.
Sukk. 14: 102–104, figs (1963). Type: Aiquile,
Habitat & Distribution: Dry inter-Andean valleys Prov. Campero, Dept. Cochabamba, Bolivia,
of the Río Mantaro and Río Apurimac drainage December 1954, Ritter 359, (erroneously as
systems, 2200–3000m. 539 in the protologue) (U).
Ecological regions: Peruvian Yungas. C. parviflorus var. comarapanus Ritter, Kakt.
Occurrence: PE (AP, AY, HV). Map 7. Südamer. 2: 683–684 (1980). Type: “east of
Comarapa”, Prov. Caballero, Dept. Santa
A very distinctive species since it is the only Cruz, Bolivia, 1954, Ritter 358 (U).
one with long, down-curved greenish-white C. parviflorus var. herzogianus (Backeb.) Backeb.,
flowers. However, there are some forms which Descr. Cact. Nov. 3: 5 (1963). Basionym: C.
produce a pink or reddish colouration of the herzogianus Backeb.
hypanthium at anthesis and it is then possible to Echinopsis parviflora (K. Schum.) Anceschi &
confuse them with C. pungens Ritter. In these Magli, Cactusinhabitat South America
cases the thick grey-green stems compared to the 2011/2013 p.39 (2013). Basionym: Cereus
thin dark green stems of C. pungens may be a parviflorus K. Schum.
useful character. As far as I am aware it is the A shrub to 3m tall branching from the base
only species which appears to be self-fertile. and higher. Stems spreading, ±3cm diameter with
It is rather surprising that this species and C. 12–14 ribs bearing close-set areoles. Spines
pungens Ritt. are the only two taxa occurring in differentiated; radials 5–9, 3–7mm long, dark
Peru. Even more so since they are highly disjunct yellow; centrals at least 1, longer and thicker.
being separated from the nearest of the remaining Flowers 2.5–3.5cm × 8mm, hypanthium pale
species by a distance of at least 450km. The yellow at first becoming red, perianth
nearest Bolivian species is C. viridiflorus Backeb., unexpanded; inner tepals purple. Fruit globose,
found in humid valleys north-west of La Paz. ±1cm diameter, yellow (Figures 27 & 28).
Ecologically, the two areas are relatively similar
and both are termed “Yungas” by Olsen et al. Etymology: from the Latin parvus and flos;
(2001). meaning “small-flowered”, the character which

Figure 29. Cleistocactus pungens LM0363.02 Figure 30. Cleistocactus reae LM0498.01 north of
west of Chincheros, Peru, 2060m. Quime, Bolivia, 2760m.
Schumann considered to distinguish it from the basitonically branching, thinner-stemmed plant
other species he knew. with magenta flowers. On occasion, however, I
suspect they produce hybrids which, although
Habitat & Distribution: Seasonally dry inter having intermediate characters, can be difficult to
Andean valleys amongst shrubs and trees in stony recognise as such. A remarkable example is plants
ground, 1300–3000m. found south east of Pasorapa, where the tall
Ecological regions: Bolivian Montane Dry Forest. yellow-spined plants have large magenta flowers
Occurrence: BO (CB, CH, SC). Map 5. copiously covered in long white hairs.
As one of the oldest names in the genus and a 15. Cleistocactus pungens Ritter, Taxon 13: 115
species with a large distribution it has acquired (1964). Type: Río Pampas, frontier of Depts
several synonyms. The defining characters of the Ayacucho and Apurímac, Peru, 1957, Ritter 664
species are its mesotonic branching habit, few (U).
spines and short red flowers. This combination is C. brevispinus Ritter, Kakt. Südamer. 4: 1360–
not found in the other species with which it is 1361, fig. (1981). Type: Pachachaca gorge near
often sympatric. In the west of its distribution it Abancay, Dept. Apurimac, Peru, 1964, Ritter
occasionally grows in close proximity to C. 1297 (U).
buchtienii Backeb., although the latter prefers A short shrub to 1.5m with stems branching
more exposed situations. Both species can form from the base. Stems grey-green, 2.5–3.5cm
large mesotonically branched shrubs, but C. diameter with 12–15 ribs, the areoles 5–7mm
parviflorus can be distinguished by its fewer, apart. Spines reddish brown; radials 8–12, 2–
stiffer spines and shorter, red flowers. In the east 10mm long; centrals 2–5, 2–5cm, stronger.
its range overlaps extensively with that of C. Flowers 7–8cm × 6–8mm diameter, red. Fruit
candelilla Cárd. and the two can frequently be globose, ±2cm diameter, pale red (Figure 29).
found together. Generally it is easy to tell them
apart since C. candelilla is a much smaller, Etymology: from the Latin pungo, to prick. Ritter

gave no explanation but presumably, referring to
the sharp spines.
Habitat & Distribution: Seasonally dry inter

Andean valleys of central eastern Peru, 1100–
2400m.
Ecological regions: Peruvian Yungas.
Occurrence: PE (AP, AY, HV). Map 7.
Only this and one other species, C.

morawetzianus Backebg., are found in Peru. Both
species grow in the same region, along the valleys
of the lower Río Apurimac and its tributaries, and
although occasionally found sympatric they
generally occur at different altitudes with C.
pungens preferring the lower slopes below
2400 m. When found together they are relatively
easy to distinguish, especially when in flower.
16. Cleistocactus reae Cárd., Cactus (Paris) 12:

251–252, figs (1957). Type: Quime, Prov.
Inquisivi, Dept. La Paz, Bolivia, 3000m,
November 1956, Cárdenas 5490 (LIL 531554,
holo., US iso.).
A many stemmed shrub branching from the

base. Stems erect, 1–2m × 3–4.5cm diameter with
18–20 ribs. Areoles only 5mm apart bearing 16–20 Figure 31. Cleistocactus ritteri LM0502.02 from
spines, radials 12–16, 5–10mm long, hyaline; the type locality near La Plazuela, Bolivia, 1070m
centrals 3 or 4, 1–3cm long, stronger, yellowish. in culture. Photograph: Martin Lowry.
Flowers clustered near the stem apex, 5–7cm × 6–
9mm, straight, presented perpendicular to the 17. Cleistocactus ritteri Backeb., Kakt. and.
stem, hypanthium scales with brown hairs; tepals Sukk. 10: 163–165, fig. (1959). Type: a flowering
strawberry red. Style not exerted. Fruit globose to plant of Ritter 325 growing in the collection of Dr.
2.5cm diameter. Seeds shiny, dark brown, 1.5mm Karius at Muggensturm (Baden). Ritter gave the
long (Figure 30). details of this collection as: at the Río La Paz,
Plazuela, Prov. Sud-Yungas frontier with Prov.
Etymology: named to honour the Bolivian Loayza, Dept. La Paz, Bolivia, 1100m, 1953.
agronomist Julio Rea who worked at the Lectotype (designated here): the illustration in the
Universidad Mayor de San Simón in Cochabamba protologue.
from where Cárdenas made his descriptions of Cephalocleistocactus ritteri (Backeb.) Backeb., Die
cacti. Cactaceae VI: 3691, figs. (1962).
A few-stemmed shrub, 1–2m high, branching
Habitat & Distribution: Amongst grass and from the base. Stems ascending, occasionally
shrubs in loamy soil on steep slopes of clearings prostrate, thin, ±3cm diameter, with 12–14 ribs
in the Yungas, 2700–3000m. bearing areoles 5mm apart. Radial spines to 30,
Ecological regions: Bolivian Yungas. 6–8mm long, very thin and fragile, white. Central
Occurrence: BO (LP). Map 6. spines – when distinguishable – a little longer and
thicker, pale yellow. Flowers from areoles near
One of several micro-endemic species. It has the stem apices producing additional conspicuous
some similarities with the typical form of C. white bristles, 3–4cm × 5mm, lemon yellow–
buchtienii and grows in an area only 40km further orange. Fruit green, 9–12mm diameter. Seeds
west at a similar altitude. At present it is little black, pyriform, 0.8mm × mm (Figure 31).
known and further observations are required to
determine its status. Etymology: named to honour Friedrich Ritter,
German explorer who spent many years searching
for cacti in South America and was the first to
discover the species.

Figure 32. Cleistocactus samaipatanus GC854.01 Figure 33. Cleistocactus samaipatanus GC918.04
west of Ipati Jctn, Santa Cruz, 1014m from north of Vallegrande, Bolivia in culture.
Photograph: Graham Charles Photograph: Graham Charles
Habitat & Distribution. On slopes beneath trees undoubtedly of the correct material but possibly
in humid Yungas forest of the Río La Paz not available when Backeberg made his
drainage, 1000–1100m. Currently known only description since he did not publish it in the
from this one small area. protologue using instead a photograph by Kilian.
Ecological regions: Bolivian Yungas. A second photograph published by Backeberg in
Occurrence: BO (LP). Map 9. Die Cactaceae (Abb. 3352) is clearly of the same
plant as in the protologue but taken from a
There is some debate regarding the validity of different angle. In view of this situation I have
this name since Backeberg explicitly cited the chosen to select the illustration in the protologue
type as a living plant in the collection of Dr Karius as the lectotype for this noteworthy species.
of Muggenstürm, Baden thus rendering his A third illustration (Die Cactaceae II: 1023
description invalid according to ICN Art 8.4. Abb. 956) was published prior to the description
However, according to Eggli et al. (1995), there where the text repeats Ritter’s description, in his
had been multiple specimens of Ritter 325 present 1956 catalogue, of the flower colour as orange.
at ZSS since 1954 thus, in their opinion, This colour has never been seen on plants in
validating the name. Undoubtedly though, this cultivation and was not found by the author on
material was not consulted by Backeberg. Ritter’s plants at the type locality. A red or pink flowered
subsequent re-description in 1980 (Kakt. form sometimes encountered in cultivation is said
Südamer. 2: 687–688) cannot be considered a to be a hybrid of which the other parent is
validation since he did not specify a replacement unknown.
type even though he had deposited material of the
type collection at ZSS. His illustration (Abb. 659) 18. Cleistocactus samaipatanus (Cárd.) D.
was actually a reprinting of Backeberg’s Abb. Hunt, Bradleya 5: 92 (1987). Basionym:
3353 from Die Cactaceae, presumably a Bolivicereus samaipatanus Cárd.
photograph of the living plant in the collection of Bolivicereus brevicaulis Ritter, Kakt. Südamer. 2:
Dr Karius. This photograph (Abb. 3353) is 703, figs. (1980). Type: Mairana, Prov. Florida,

Dept. Santa Cruz, Bolivia, 1958, Ritter 839, north-eastern Andes, 1300–2100m.
(U). Ecological regions: Southern Andean Yungas,
B. croceus Ritter, Kakt. Südamer. 2: 704–705, fig. Bolivian Montane Dry Forest, Dry Chaco.
(1980). Type: “probably” Puente Azero, border Occurrence: BO (CH, SC). Map 4.
of Prov. Tomina and Prov. Azero, Dept.
Chuquisaca, Bolivia, Ritter 1471, (U). Cárdenas considered his Bolivicereus
B. rufus Ritter, Kakt. Südamer. 2: 703–704, fig. samaipatanus as distinct based solely on the wide
(1980). Type: Puente Azero, border of Prov. opening of the perianth compared with the then
Tomina and Prov. Azero, Dept. Chuquisaca, known members of the genus Cleistocactus. It is
Bolivia; Ritter 840, (U). now one of three taxa with flowers having
B. samaipatanus Cárd., Cact. Succ. J. (US) 23: 91 expanded perianths retained in Cleistocactus due
(May 1951). Type: near “El Fuerte” de to the otherwise conforming characters,
Samaipata, Prov. Florida, Dept. Santa Cruz, particularly the lateral positioning of the flowers.
Bolivia, 1890m, May 1948; A. Corro in The species is quite widely distributed and has
Cárdenas 4395 (LIL 531550, holo.; US). been found growing sympatrically with six other
B. samaipatanus var. multiflorus Cárd., Cact. species of the genus without the appearance of
Succ. J. (US) 23: 93–94, fig. (May 1951). Type: hybrids. Notably however, the one species with
La Angostura, Prov. Florida, Dept. Santa which hybrids have been raised in cultivation,
Cruz, Bolivia, September 1949, Cárdenas Cleistocactus winteri ssp. winteri, to produce the
4395a (LIL 531549, holo.; US). cultivar ‘Jupiter’ (Mottram, 1989), is not amongst
Borzicactus samaipatanus (Cárd.) Kimnach, Cact. these six species.
Succ. J. (US) 32: 93 (1960). Basionym: The apparent disjunction in distribution in the
Bolivicereus samaipatanus Cárd. Andean portion of its range is primarily due to
B. samaipatanus var. multiflorus (Cárd.) Krainz, lack of exploration in the area between
Kakteen, Lief. 21: C Vb. (1962). Basionym: Vallegrande and Moteagudo. The mountain
Bolivicereus var. multiflorus Cárd. ranges in the region seem highly suitable for the
Echinopsis samaipata (Cárd.) Anceschi & Magli, species and it is likely to be found there
Cactusinhabitat South America 2011/2013 eventually.
p.39 (2013). Basionym: Bolivicereus
samaipatanus Cárd. b. ssp. divi-miseratus (Cárd. ex Backeb.) M.
A sprawling, decumbent or erect shrub Lowry comb. et stat. nov. Basionym:
branching from the base. Stems to 1.5m × 3–4cm Bolivicereus samaipatanus var. divi-miseratus
diameter with 14–16 low rounded ribs bearing Cárd. ex Backeb., Natl. Cact. Succ. J. 6: 9, fig.,
areoles 3–4mm apart. Spines 13–22, unequal, 4– (March 1951) nom. inval. [Art. 35.1]; Die
30mm, yellow or brown at first becoming grey. Cactaceae 2: 972, 973, (Mar 1959). Type: Fig. 892
Flowers 3.5 × 2cm, bright red; hypanthium S- of a flowering plant on p.972 of the protologue).
shaped, perianth expanded obliquely. Stamens Bolivicereus samaipatanus ssp. divi-miseratus
and style exerted with filaments and anthers (Cárd.) Guiggi, Cactology 3: 6, (2012) nom.
purplish. Nectar chamber closed by staminodial inval. (Art. 41.5). Basionym incorrectly cited
hairs. Fruit ±globose, 9–11 × 7–9mm, reddish as: B. samaipatanus var. divi-miseratus Cárd.
brown. Borzicactus samaipatanus var. divi-miseratus
(Cárd. ex Backeb.) Krainz, Kakteen, Lief. 21:
Key to subspecies of C. samaipatanus C Vb. (1962). Basionym: Bolivicereus
1. Stems sprawling or decumbent, spines few to samaipatanus var. divi-miseratus Cárd. ex
many, stiff (Andean foothills) (Figures 32 & 33) Backeb.
a. ssp. samaipatanus Etymology: referring to the place of discovery of
Stems erect, spines many, fine (Chiquitano the plant, the Serrania Divi Miserato, a range of
Dry Forest, Prov. Chiquitos) (Figures 34 & 35) hills northeast of Roboré, Bolivia.
b. ssp. divi-miseratus
Habitat & Distribution: Arid rocky outcrops and
a. ssp. samaipatanus hills amongst shrubs in the mountains east of
Roboré, 600–900m.
Etymology: referring to the place of discovery of Ecological regions: Chiquitano Dry Forest.
the plant, Samaipata, then a small market town Occurrence: BO (SC). Map 4.
in the Andean foothills west of Santa Cruz.
Long considered as just a disjunct variant this
Habitat & Distribution: On cliffs and steep slopes form has become adapted to a very different
in seasonally dry inter Andean valleys of the environment than that of the Andean foorhills. It

Figure 34. Cleistocactus samaipatanus ssp. divi-miseratus Bercht 3781, Aguas Calientes, Santa Cruz, Bo-
livia, 253m. Photograph: Ludwig Bercht.
grows erect rather than decumbent, has more ,
generally hyaline spines and longer flowers which
lack the paler edge to the tepals.
Cárdenas' original description of the plant as a

variety has to be considered invalid since it was
published two months prior to the description of
the genus and type subspecies. The name remained
so until validated by Backeberg in 1959. Cárdenas
had intended the type to be a plant he collected in
February 1950 at 900m in the Sierrania Divi
Miserato of Province Chiquitos, Dept. Santa Cuz,
Bolivia and subsequently preserved in his
herbarium as Cárdenas 4569 (LIL 531545).
19. Cleistocactus smaragdiflorus (F.A.C.

Weber) Britt. & Rose, Cact. 2: 174 (1920).
Basionym: Cereus colubrinus var. smaragdiflorus
F.A.C. Weber.
Cereus colubrinus var. smaragdiflorus F.A.C.
Weber, Dict. hort. Bois p.281 (1894). Type:
Figure 35. Cleistocactus samaipatanus ssp. divi- from cultivated material, not designated.
Neotype, selected by Hunt & Taylor, Cact.
miseratus Bercht 3781, Aguas Calientes, Santa
Syst. Init. 21: 6 (2006): Gürke, Bluh. Kakt. 2: t.
Cruz, Bolivia, 253m.
87 (1906).
Photograph: Ludwig Bercht.

Figure 36. Cleistocactus smaragdiflorus Figure 37. Cleistocactus smaragdiflorus
LM0523.01 east of Entre Rios, Bolivia, 1180m. LM0243.03 south of Alemania, Argentina, 1300m.
The form previously known as C. rojoi. Photograph: Martin Lowry.
C. smaragdiflorus var. gracilior Backeb.,
C. smaragdiflorus (F.A.C. Weber) Speg., Anal. Kakteenlexikon p.89 (1966), nom. inval.. Type:
Mus. Buenos Aires xi: 482 (1905). Basionym: not designated.
Cereus baumannii var. smaragdiflora (sic) C. smaragdiflorus fma. rojoi (Cárd.) Ritter, Kakt.
F.A.C. Weber. Südamer. 2: 678 (1980). Basionym: C. rojoi
Cleistocactus azerensis Cárd., Cact. Succ. J. (US) Cárd.
33: 74, figs., (1961). Type: Puente Pacheco, C. villamontesii Cárd., Cact. Succ. J. (US) 33: 76–
Prov. Azero, Dept. Chuquisaca, Bolivia, 77, figs., (1961). Type: Angosto de Villa
1100m, April 1959, Cárdenas 5522 (LIL, Montes, Prov. Gran Chaco, Dept. Tarija,
531534, holo.; US). Syn. nov. Bolivia, 460m, May 1958, Cárdenas 5524 (LIL
C. ferrarii Kiesling, Hickenia 2: 37, fig. (1984). 531598, holo.; US, iso.).
Type: Cerro de la Cruz, Sierras de Zapla, C. villamontesii var. longiflorior Backeb.,
Capital, Jujuy, Argentina, April 1974, Kiesling Kakteenlexikon p.90 (1966), nom. inval. Type:
1722 (SI). not designated.
C. parapetiensis Cárd., Cact. Succ. J. (US) 24: A small shrub with erect, arching or
182–183, figs., (1952). Type: on the road from decumbent stems to 1m long. Stems 2–3cm
Camiri to Charagua, Prov. Cordillera, Dept. diameter with 12–14 ribs bearing brown-felted
Santa Cruz, Bolivia, 900m, March 1949, areoles. Spines differentiated, radials 10–14, up
Cárdenas 4821 (LIL, not found; US 2906758, to 1cm long; centrals 4–6, 15–35mm long,
iso). stringer, yellowish or brown. Flowers 4–5cm ×
C. rojoi Cárd., Cact. Succ. J. (US) 28: 56, figs. ±8 mm, straight, hypanthium red with scales and
(1956). Type: between Entre Rios and Villa white bristles, perianth unexpanded, outer tepals
Montes, Prov. O’Connor, Dept. Tarija, Bolivia, yellow, inner tepals vivid emerald green. Fruit
600m, March 1952, Cárdenas 5052 (LIL, globose, ±1.5cm diameter (Figures 36 & 37).
531576, holo.; US, iso.).
C. smaragdiflorus var. flavispinus Borg, Cacti Etymology: from the Latin smaragdus meaning
p.151, (1937). Type: not designated. “emerald” and referring to the bright green tepals.

Figure 38. Cleistocactus strausii LM s.n. north of Santa Ana, Bolivia, 2200m.
Habitat & Distribution: Amongst trees and
bushes in the seasonally dry hills and valleys in
the eastern foothills of the Andes, 500–1500m.
Ecological regions: Southern Andean Yungas and
Dry Chaco.
Occurrence: AR (CT, JY?, SA, TM); BO (CH, SC,
TR). Map 8.
Long considerd to be an Argentinian endemic

but now taken to include several forms (C.
azerensis, parapetiensis, rojoi and villamontesii)
in Bolivia described by Cárdenas. These forms
differ only marginally from the type in that their
stems tend to be thinner and their central spines
shorter. Their flowers are more or less identical to
the type form with the exception that the green
inner tepals may be shorter and fewer giving
them a slightly less contrasting colouration.
It can be found growing sympatrically with C.
brookeae Cárd. (C. vulpis-cauda Ritter &
Cullmann) in Bolivia and with C. baumannii
(Lem.) Lem. in Argentina and Bolivia.
It is worth noting that Backeberg’s illustration
Figure 39. Cleistocactus strausii Preston-
(Die Cactaceae VI: 3694 Abb. 3354) of what he
Mafham 189 from east of the Condor Pass, Tarija,
thought might be C. parapetiensis Cárd. is in fact
Bolivia in culture. Photograph: Graham Charles.

what he later described as Cephalocleistocactus In habitat it can occur sympatrically with both
schattatianus! C. micropetalus and C. hyalacanthus, either
singly or together, with no hybrids between them.
20. Cleistocactus strausii (Heese) Backeb., This clearly demonstrates their adaptation to
Kakteen-Freunde 3: 121 (1934). Basionym: pollination by distinct species of hummingbird.
Pilocereus strausii Heese. Like Darwin’s prediction for the orchid
Borzicactus straussii (Heese) Berger, Entwickl. Angraecum sesquipedale Thouars one could
Kakt. 53 (1926). Basionym: Pilocereus strausii postulate that there is a species of hummingbird
Heese. (rather than hawkmoth) with a beak sufficiently
Cephalocereus straussii (Heese) Houghton, Cact. long to reach the nectaries of C. strausii, which
Succ. J. (US) 1: 170 (1930). Basionym: has the longest flower of the three cacti, yet
Pilocereus strausii Heese. cannot enter the curved flowers of C.
Cereus straussii (Heese) Vaupel, Monatschr. Kakt. hyalacanthus nor the tightly closed flowers of C.
23: 37 (1913). Basionym: Pilocereus strausii micropetalus. Presumably there are two other
Heese. hummingbird species with beak characteristics
Demnoza straussii (Heese) Frič, Möller’s Deutsche allowing them access to the nectar in these other
Gärtn.-Zeit. 44: 170. (1929). Basionym: plants.
Pilocereus strausii Heese. Cleistocactus strausii has significant horticul-
Echinopsis nothostrausii Anceschi & Magli, tural value and is often grown outdoors in warm
Cactusinhabitat South America 2011/2013 climates where it can produce huge specimens. It
p.39 (2013). Replaced synonym: Pilocereus has also been hybridized with species from other
strausii Heese. (Nom. illeg. [Art. 52.1]). genera, notably Echinopsis to produce flowers
Pilocereus strausii Heese, Gartenflora 56: 410, fig. that open more widely but retain the colour of C.
49, (1907). Type: from cultivated material, strausii.
hort. Berlin, Bolivia, not preserved. Lectotype,
selected by Hunt & Taylor, Cact. Syst. Init. 21: 21. Cleistocactus tominensis (Weingart)
6 (2006): the fig. cited. Backeb., Kaktus-ABC p.190 (1936). Basionym:
A few-stemmed shrub 1–3m tall branching Cereus tominensis Weingart.
from the base. Stems stout, 4–8cm diameter with Borzicactus tominensis (Weingart) Borg, Cacti
25–30 ribs bearing close-set areoles. Spines p.147 (1937). Basionym: Cereus tominensis
differentiated, centrals, about 4, to 2cm, acicular, Weingart.
pale yellow; radials, 30–40, 15–20mm, very fine Cereus tominensis Weingart, Monatschr. Deutsch.
like hairs, pure white. Flowers from upper third of Kakteen-Ges. 3: 117–118 (1931). Type: dry veg-
stem, horizontal, 8–9cm × 7–9mm, dark red- etation in the valley of Tomina, Prov. Tomina,
purple, hypanthium covered with silky brown Dept. Chuquisaca, 1800–2000 m, 1926–28,
hairs, perianth unexpanded with long-exserted Troll 174, not preserved.
style. Fruit globose, about 2cm diameter, reddish- C. mendozae Cárd., Cact. Succ. J. (US) 35: 202,
brown with brown hairs (Figures 38 & 39). figs., (1963). Type: between Tomina and
Padilla, Prov. Tomina, Dept. Chuquisaca, Bo-
Etymology: named to honour Leopold Straus, a livia, 2200m, March 1958, Cárdenas 5550,
German merchant from Bruchsal, cactus hobbyist (LIL, not found; US, not recorded). Lectotype
and co-founder of the German Cactus Society (designated here): the photograph with the
(DKG). protologue.
Echinopsis tominensis (Weigart) Anceschi &
Habitat & Distribution: Steep cliffs and mountain Magli, Cactusinhabitat South America
sides of the seasonally dry valleys of central 2011/2013 p.40 (2013). Basionym: Cereus tomi-
Province Tarija, 1700–2700m. nensis Weingart.
Ecological regions: Central Andean Puna and An erect shrub to 1–2m tall branching from
margins of Southern Andean Yungas. the base. Stems ±5 cm diameter with 13–22 low,
Occurrence: BO (TR). Map 5. transversely-furrowed ribs. Areoles brown-felted,
about 1 cm apart bearing 1–20 acicular spines;
Probably the most easily recognised species of centrals 1–3, 10–15mm long, yellow; radials 0–20,
the genus although it spent time in several genera shorter and finer. Flowers produced in profusion
before its recognition as a Cleistocactus by from upper part of stem, straight, 2.5–3cm × 7–
Backeberg in 1934. 9mm, hypanthium at first green becoming red at
anthesis, perianth unexpanded, tepals green.

Figure 40. Cleistocactus tominensis LM0963A.01 Figure 41. Cleistocactus tominensis
south of Rendencion Pampa, Bolivia, 2260m. The BCMdV200.01 from north of Nuevo Mundo, Bo-
form previously known as C. mendozae. livia, 1250m in culture.
Figure 42. Cleistocactus variispinus LM0197.01 Figure 43. Cleistocactus viridiflorus LM0444.01
from north of Sorata, Bolivia, 3060m in culture. near Acupata, Bolivia, 3140m.

Style slightly exserted. Fruit globose, about 15mm style not exserted. Fruit globose, 1.5–2cm diame-
diameter, pale pink–brown (Figures 40 & 41). ter, violet-red (Figure 42).
Etymology: referring to the place of first discovery Etymology: from the Latin varius and spinus re-
of the species, Tomina, a small town in the eastern ferring to the spines being of variable length. The
part of Department Chuquisaca, Bolivia. apparent correction used in the NCL is copied from
Eggli et al. (1995) where Ritter’s spelling was, in
Habitat & Distribution: Deciduous shrub lands error, considered a misprint (ICN Rec. 60G.1).
and forest of the seasonally dry inter-Andean val-
leys along the Río Grande drainage, 1000–3100m. Habitat & Distribution: Valleys and slopes of
Ecological regions: Bolivian Montane Dry Forest humid mountain forests above the Yungas, 3000 –
and margins of Southern Andean Yungas. 3600m.
Occurrence: BO (CB, CH, PO, SC). Map 4. Ecological regions: Central Andean Wet Puna and
Bolivian Montane Dry Forest.
In the absence of a preserved type for this Occurrence: BO (LP). Map 9.
species it is unfortunate that the illustration in
the protologue is of a small seedling that is not at One of several species which produces addi-
all representative of the mature plant. tional long spines and bristles on areoles in the
The most striking character of this species is flowering zone of the stem to produce a pseudo-
the colour-changing flower. The buds start bright cephalium. The production of this pseudo-cephal-
green and remain so until anthesis at which time, ium was the reason why Backeberg either
presumably to indicate readiness for pollination, transferred to, or described new names under Rit-
the hypanthium turns bright red. The effect is ter’s genus Cephalocleistocactus. Ritter
particularly marked in the form Cárdenas called adamantly disapproved of these names since for
C. mendozae and in the flowering season these him the pseudocephalia of these plants were not of
plants can be seen with many flowers at all stages the same form as those produced by his Cephalo-
of development giving the impression of multi- cleistocactus chrysocephalus.
coloured flowers as noted in Weingart’s first de- Ritter’s illustration (Kakt. Südamer. 2: Abb.
scription. 677) shows a large sprawling mass of stems on a
The species is quite variable with plants from hillside apparently bare of trees. The stems ap-
the bottom of the Río Grande valley having far pear to be quite short and thick, unlike those of
fewer, sometimes only one stem and often a com- plants currently in cultivation under this name
plete absence of radial spines (Figure 41). The which are similar to the plant described by Backe-
flowers of this form are, however, similar to the berg as Cephalocleistocactus schattatianus. Ritter
type and behave in an identical manner. The de- also quotes the habitat altitude as 2000m which is
scription given above is amplified from the origi- considerably below that at which I have seen
nal to reflect these differences. plants similar to those in cultivation. Ritter also
goes to great length to argue that Backeberg’s C.
22. Cleistocactus variispinus Ritter, Taxon 13: schattatianus may not be the same as C. variispi-
114 (1964); Kakt. Südamer. 2: 689–691, figs., nus. Thus there is still some confusion over the
(1980). Type: Mina Cascabel, Prov. Lareca, Dept. correct application of this name that would bene-
La Paz, Bolivia, 2000m, 1953, Ritter 108 (U fit from a careful exploration of the area around
117850B). the type locality.
Cephalocleistocactus schattatianus Backeb.,
Descr. Cact. Nov. III: 5 (1963), nom. inval. [Art 23. Cleistocactus viridiflorus Backeb., Descr.
40.1]. Cact. Nov. III: 5 (1963). Type: a plant of Ritter 323
A many-stemmed erect or decumbent shrub in the collection of Saint-Pie. Ritter gives details
freely branching from the base and lower half. for this number as: Ayata, Prov. Muñecas, Dept.
Stems 0.5–1 (-2)m × 3–5cm diameter with 14–18 La Paz, Bolivia, 3300m, Oct 1954., Lectotype, des-
ribs bearing areoles only 3–4mm apart. Spines ignated by Backeberg: Kakteenlexicon p.90, fig. 61
30–40, pale yellow to brownish-yellow, most < (left) (1966).
10mm but occasionally 1 or 2 longer, 2–4cm. Flow- Cleistocactus palhuayensis Ritter & Shahori,
ers from upper part of stem from areoles produc- Kakt. Südamer. 2: 688, figs., (1980). Type:
ing additional bristles and spines, 3.5–4.5cm × lower Río Palhuaya, Prov. Muñecas, Dept. La
6–8 mm, hypanthium crimson red–orange, tepals Paz, Bolivia, 1954, Ritter 324 (U, not found;
white with pale green tips, spreading slightly, ZSS T4911 & T4912, iso.)

C. palhuayensis var. camachoensis Ritter & Sha- claims that this species grows at the highest ele-
hori, Kakt. Südamer. 2: 688–689, figs. (1980). vation of all cleistocacti, where in winter the tem-
Type: upper Río Palhuaya, Prov. Camacho, perature sinks below freezing at night.
Dept. La Paz, Bolivia, 3200m, Ritter s.n. (U,
not found). Note: Eggli (Englera 16: 401 24. Cleistocactus winteri D. Hunt, Bradleya 6:
(1995)) gives Ritter 835 as the probable type 100 (1988). Replaced synonym: Winteria au-
number for this taxon, however, the location reispina Ritter.
details are different from those given in the Borzicactus auriespinus (Ritter) Rowley, Repert.
protologue. Also Ritter clearly states there Pl. Succ. XXIV - 1973: 6 (1975), nom. inval.
that this taxon has no number, however he [Art. 41.5].
later (Kakt. Südamer. 4: 1637) assigns this Cleistocactus auriespinus (Ritter) D. Hunt (non C.
number. auriespinus Frič), Bradleya 5: 92 (1987), nom.
Cephalocleistocactus pallidus Backeb., Kakteen- illeg. [Art 41].
lexicon p.79, fig. (1966). Type: Palhuaya gorge, Hildewintera auriespina (Ritter) Ritter, Kakt.
Consata basin, Dept. La Paz, Bolivia, Ritter and. Sukk. 17: 11 (1966), nom. inval. [Art.
324. 35.1, 41.5]. Intended replaced synonym: Win-
An erect shrub, branching from the base. teria auriespina Ritter.
Stems 1–1.6m × 2–4cm diameter with about 12– Hildewintera auriespina (Ritter) Ritter ex G.D.
17 low ribs. Areoles with tawny felt and 15–20 yel- Rowley, Repert. Pl. Succ. XVII in Regnum Veg.
lowish spines, radials 15–18, < 1cm long, thin; 54: 15. (1968), nom. illeg. [Art 52.1].
centrals 1 or 2, to 2cm long, thicker. Flowers al- Loxanthocereus auriespinus (Ritter) Buxbaum,
most apical, about 3.5 cm × 6–8mm diameter from Kakteen (H. Krainz) 57: CVb (1974), nom.
areoles producing additional longer spines, hy- inval. [Art. 41.5].
panthium with short brown hairs, basally reddish, Winteria auriespina Ritter, Kakt. and. Sukk. 13:
upper half pale yellow–green, tepals pale yellow– 4–8, figs. (1962), nom. incorr. [Art 11.4]. Type:
green, barely expanded. Style not exserted. Fruit Yapacani gorge on overhanging rocks, Agua
globose, about 1.5cm diameter, reddish with Clara, Prov. Florida, Dept. Santa Cruz, Bo-
brown hairs (Figure 43). livia, July 1958, Ritter 846 (U 097816B).
Winterocereus auriespinus (Ritter) Backeb.,
Etymology: from the Latin viridus and flos; Kakteenlexikon p.455 (1966).
“green-flowered”. A procumbent or pendant shrub with stems
branching from the base. Stems 1–2m × 3–7cm
Habitat & Distribution: Amongst trees and shrubs with 14–19 ribs bearing closely set areoles. Spines
on slopes of humid valleys in the northern Yun- 15–30, variable, golden yellow or white, 5–50mm
gas, 1400–3200m. long. Flowers 4–6 × 5cm diameter, orange or red,
Ecological regions: Bolivian Yungas and Bolivian perianth expanded and nearly regular. Outer
Montane Dry Forest. tepals linear or spathulate, 2–3.5cm × 3–5mm,
Occurrence: BO (LP). Map 9. spreading; inner tepals spathulate, white–pale-
pink, 3–8mm × 3–5mm, incurved against the
Previously referred, in the NCL, to C. filaments. Stamens and style exserted, filaments
buchtienii Backeb., this species is not only highly concolourous with outer tepals, anthers orange or
disjunct but grows in a very different environ- magenta. Fruit globose, 7–10mm diameter,
mental niche and has much shorter green rather reddish green.
than purple flowers.
This species has several characters in common Habitat & Distribution: Sandstone cliffs in semi-
with the previous taxon which grows in a similar deciduous humid forest of the north-eastern
environment nearby and they may only be ecolog- Andean valleys, 1300–1400 m.
ical variants of one species. They differ primarily Ecological regions: Bolivian Montane Dry Forest.
in growth form and branching pattern. Cleisto- Occurrence: BO (SC). Map 2.
cactus variispinus regularly branches mesotoni-
cally and can eventually form a small tree The nomenclatural history of this species is
whereas this habit has not been seen in the cur- highly confusing and even propagated into
rent species. I hesitate to synonymise them here confusion over the names of both subspecies.
since I have not observed plants of this species or Unravelling the confusion took several attempts
its synonyms at their respective type localities. over a number of years and I do not propose to
In his 1956 catalogue (Winter, 1956) Ritter revisit the issues here simply referring the

Figure 45. Cleistocactus winteri ssp. winteri
LM0021.01 from west of Mairana, Bolivia, 1430m
in culture. Photograph: Martin Lowry.
Figure 44. Cleistocactus winteri ssp. winteri

LM0021.01 at its only locality west of Mairana,
Bolivia, 1430m. Photograph: Martin Lowry.
Figure 47. Cleistocactus winteri subsp. colade-

Figure 46. Cleistocactus winteri ssp. colademono mono Krahn 950 ( Type collection) from Cerro El
LM0587.03 on Cerro El Fraile, Dept. Santa Cruz, Fraile, Dept. Santa Cruz, Bolivia, in culture.
Bolivia, 1350m. Photograph: Martin Lowry. Photograph: Graham Charles.

interested reader to two excellent papers by a feature also present in C. brookeae.
Kiesling & Metzing (2004) and Metzing & Even after nearly sixty years the taxon is still
Kiesling (2007). As these authors say, the only known from a single location where the old
confusion was not caused by any neglect of the road from Santa Cruz to Cochabamba runs
rules and recommendations “but rather, it shows alongside the Río Yapacani north of Pampa
the complexity of the Code ... and its many Grande, an area of about 0.06km2. I have been
pitfalls”. Essentially, strict application of the Code observing the plants there for 20 years and
(McNeill et al., 2012) does not allow retention of although the road has been much developed in
the generic name Hildewintera since Ritter did that time there are still many specimens visible
not explicitly state the page number of his earlier on the rocks.
description of the genus Winteria. Thus, if one
wishes to consider these plants as distinct from b. ssp. colademono D. Hunt, Cact. Syst. Init. 20:
Cleistocactus, the earliest validly published 14 (2005). Replaced synonym: Hildewintera
generic name is Winterocereus Backeb., which is colademononis Diers & Krahn.
where Metzing and Kiesling place the two Hildewintera colademononis Diers & Krahn,
subspecies as separate species. The taxa are Kakt. and. Sukk. 54: 221, figs (31 July 2003).
retained here in Cleistocactus based on their Type: Cerro el Fraile, Prov. Florida, Dept.
many shared features including stem, flower, fruit Santa Cruz, Bolivia, 2000; Krahn 950
and seed characteristics, and pending further (KOELN, holo.).
molecular studies on the circumscription of the H. polonica E. Foik & V. Foik, Cactaceae etc.
genus. Their few differing floral traits, although (Bratislava) 13: 68–70 (15 July 2003), nom.
considerable, are taken to be derived from the inval. [Art. 40.7].
basic floral structure of Cleistocactus. H. polonica E. Foik & V. Foik, Cactaceae etc.
(Bratislava) 14: 43–50, (2004), nom. inval.
Key to subspecies of C. winteri [Art. 38.13 & 41.5].
(Adapted from Kiesling & Metzing, 2004) Borzicactus colademononis (Diers & Krahn) Lodé,
1. Stems almost rigid, erect when young, Cact.-Avent. Int. 97: 2 (2013, published 28-12-
pendulous later, spines golden yellow, 2012). Basionym: Hildewintera colademononis
hypanthium curved without axial spur, nectar Diers & Krahn.
chamber closed by diaphragm, outer tepals orange Cleistocactus colademononis (Diers & Krahn)
(Figures 44 & 45) a. ssp. winteri Mottram, CactusWorld 24: 21 (2006).
Stems flexible, always pendulous, spines like Basionym: Hildewintera colademononis Diers
hair, white, hypanthium geniculate with axial & Krahn.
spur, no diaphragm, outer tepals red (Figures 46 Winterocereus colademononis (Diers & Krahn)
& 47) b. ssp. colademono Metzing & R. Kiesling, Taxon 56: 227 (2007).
Basionym: Hildewintera colademononis Diers
a. ssp. winteri & Krahn.
Etymology: apparently referring to the husband Etymology: the local vernacular name of the plant
of Frau Hildegard Winter from whose nursery around Samaipata, deriving from Spanish and
near Frankfurt she distributed the seeds Ritter meaning “tail of the monkey”, an allusion to its
collected in South America (Mottram, 2006). many soft white spines.
Nomenclatural issues aside, C. winteri is a The discovery of this plant in 2000–2003

remarkable plant showing features highly created a sensation in cactus horticulture and the
adapted to growth on cliffs in an occasionally very plant has rapidly spread through collections
wet habitat where for six consecutive months the around the world. Its flowers are perhaps its most
rainfall can exceed 80mm per month. The long appealing attribute. They have characters
downward pointed spines allow this rainfall and combining those of both C. winteri ssp. winteri
run-off to quickly drain from the plant. Further and of C. brookeae, leading to the suggestion that
adaptations are evident in the floral structure the plant might be of hybrid origin (Kiesling &
where the reduced and in-curved inner tepals Metzing, 2004; Mottram, 2006; Charles, 2011).
protect the nectar chamber from unwanted insect The stems also resemble those of the former C.
visitors. In addition, in the current subspecies, the vulpis-cauda Ritter & Cullmann (now considered
nectar chamber is further protected by a synonymous with C. brookeae Cárd.), adding
diaphragm created by the connate stamen bases, further weight to this suggestion. However,

Undescribed species
Although the area of distribution of the genus
is relatively well explored there are still some
areas that would benefit from further exploration.
In the past this has been difficult due to limited
access and poor road infrastructure, a situation
which is improving rapidly, particularly in
Bolivia. One particular area which has already
revealed an interesting, potentially new species,
is the Bolivian Yungas. Plants discovered in this
area have now matured in cultivation and
produced bright orange flowers (Figure 48).
Considering the disjunction in distribution of the
genus between the Peruvian and Bolivian species
it would interesting to explore this region in
greater detail. Further tantalizing photographic
images taken north-east of Independencia, north-
east of Vallegrande and east of Pulquina also
show plants which are difficult to place within any
of the species recognised here. It seems then that
we don’t yet know the complete picture for this
fascinating genus.
Distribution Maps
Figure 48. Cleistocactus sp. An undescribed

species from the Yungas east of La Paz, Bolivia.
Photograph: Graham Charles.
considering the features of all three taxa it is
potentially possible that C. winteri ssp.
colademono could be a contributor rather than a
recipient of genetic information and that C.
winteri ssp. winteri is actually the hybrid.
One of the main characters which led Kiesling
and Metzing (2004) to consider the current
subspecies as a separate species is its possession
of an axial spur on the hypanthium. They
considered this structure, sometimes associated
with sphingophily, to be, in this case, an
adaptation to ornithophily allowing humming-
birds to obtain nectar without damaging the lower
gynoecium. Although this structure is clearly
evident in the figures of their paper and they
considered it unique, careful examination of the
sectioned geniculate flower of C. wendlandiorum
illustrated by Backeberg show an identical
structure, more clearly visualized in an image by Map 1. (Central South America). Occurrence lo-
Charles (2011). Externally the flowers of C. cations for Cleistocactus baumannii (red dots) and
vulpis-cauda Ritter & Cullmann show a its subspecies horstii (green dot). In this, and sub-
pronounced bulge at the knee and a section sequent maps, shading represents elevation in
reveals that this taxon too possess a rudimentary meters. Cities with representative elevations for
axial spur. These observations give considerably this map are: La Paz: 3650m, Santa Cruz: 420m,
more weight to the argument that the two Filadelphia: 140m, Asuncion: 60m, Cordoba:
subspecies of C. winteri D. Hunt should remain 400m and Salta: 1200m.
within Cleistocactus.

Map 3. (South-western Bolivia). Occurrence loca-
tions for Cleistocactus buchtienii (northern, type
Map 2.(Eastern Bolivian Andes). Occurrence loca- form: red dots, southern, “tupizensis” form: brown
tions for Cleistocactus brookeae (yellow dots) and dots), C. candelilla (magenta dots) and C. mi-
the two subspecies of C. winteri: ssp. winteri (green cropetalus (green dots). Representative elevations
dot), ssp. colademono (red dot). Representative el- are Cochabamba: 2570m, Santa Cruz: 420m,
evations are Cochabamba: 2570m, Santa Cruz: Sucre: 2820m and Tarija: 1900m.
420m and Tarija: 1900m.
Map 4. (South-eastern Bolivia). Occurrence loca- Map 5. (Southern Bolivia). Occurrence locations
tions for Cleistocactus capadalensis (red dots), C. for Cleistocactus parviflorus (green dots), C. de-
tominensis (yellow dots), C. samaipatanus (green pendens (red dots), C. hildagardiae (yellow dots)
dots) and the approximated type locality of its sub- and C. strausii (purple dots). Representative ele-
species divi-miseratus (purple dot). Representative vations are Cochabamba: 2570m, Santa Cruz:
elevations are Cochabamba: 2570m, Santa Cruz: 420m, Sucre: 2820m and Tarija: 1900m.
420m, Sucre: 2820m and Tarija: 1900m.

Map 6. (Central west Bolivia). Occurrence loca- Map 7. (Central Peru). Occurrence locations for
tions for Cleistocactus laniceps (red dots), C. lurib- Cleistocactus pungens (red dots) and C. morawet-
ayensis (green dots) and C. reae (purple dots). zianus (green dots). Representative elevations are
Representative elevations are La Paz: 3660m and Huancavelica: 3720m, Ayacucho: 2730m and Aban-
Cochabamba: 2560m. cay: 2475m.
Map 8. (Central Andes). Occurrence locations for Map 9. (Bolivian Yungas). Occurrence locations for
Cleistocactus hyalacanthus (red dots) and C. Cleistocactus chrysocephalus (red dots), C. viridi-
smaragdiflorus (green dots). Representative eleva- florus (green dots), C. variispinus (purple dots) and
tions are Sucre: 2820m, Tarija: 1900m, Jujuy: C. ritteri (yellow dots). Representative elevations
2175m, Salta: 1200m and Tucuman: 425m. are La Paz: 3660m and Cochabamba: 2560m.

Acknowledgements
The preparation of this synopsis would not HERNáNDEZ-HERNáNDEZ, T., HERNáNDEZ, H.M.,
have been possible without the help of many DE-NOVA, J.A., PUENTE, R., EGUIARTE, L.E. &
people. In particular I am grateful to Graham MAGALLóN, S. (2011). Phylogenetic
Charles, Paul Hoxey, Mats Winberg and Holger relationships and evolution of growth form in
Kelsch for providing additional occurrence data Cactaceae (Caryophyllales, Eudicotyledoneae).
and to Graham Charles, Andreas Hofacker, Am. J. Bot. 98: 44–61.
Zlatko Janeba and Paul Laney for providing HUNT, D. & TAYLOR, N.P. (1987). New and
copies of older literature. Ludwig Bercht, Graham unfamiliar names of Cactaceae to be used in
Charles, Holger Kelsch and Paulo Robson de the European Garden Flora. Bradleya 5: 91–
Souza also provided additional images. I also 94.
thank Roy Mottram and Nigel Taylor for HUNT, D. (1999). CITES Cactaceae Checklist. RBG
enlightening feedback on nomenclatural Kew. London.
complexities. It is also a pleasure to thank all HUNT, D. (2003). Notulae Systematicae Lexicon
those friends who have travelled with me in the Cactacearum Spectantes III: Cleistocactus.
field and helped make the collection of the data Cact. Syst. Init. 15: 7–8.
such a rewarding experience. HUNT, D., TAYLOR N.P. & CHARLES, G. (2006). The
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BUxBAUM, F. (1958). The phylogenetic division of Cousin, Paris.
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Cactus Explorer 1: 13–15. Peru. Bot. Jahrb. Syst. 124: 13–30.
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Ritter Collections. Englera 16. 12–16.
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Springer. Berlin & Heidelberg. MCNEILL, J., BARRIE, F.R., BUCK, W.R., DEMOULIN,
EGGLI, U. & LEUENBERGER, B.E. (2005). The V., GREUTER, W., HAWKSWORTH, D.L.,
Cárdenas Type Specimens of Cactaceae HERENDEEN, P.S., KNAPP, S., MARHOLD, K.,
Names in the Herbarium of the Instituto PRADO, J., PRUD’HOMME VAN REINE, W.F.,
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277–281.

METZING, D. & KIESLING, R. (2007) Winterocereus colademononis 24b
(Cactaceae) is the correct name for janae 8
Hildewintera. Taxon 56: 226–228. laniceps 10
MOTTRAM, R. (1989). New Cleistocactus cultivars. samaipatanus 18a
Cact. Succ. J. (Los Angeles) 61: 154–157. samaipatanus var. divi-miseratus 18b
MOTTRAM, R. (2002). Notulae Systematicae samaipatanus var. multiflorus 18a
Lexicon Cactacearum Spectantes I: strausii 20
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Cactus World 24: 19–22. strausii 20
OLSON, D.M., DINERSTEIN, E., WIKRAMANAYAKE, Cephalocleistocactus
E.D., BURGESS, N.D., POWELL, G.V.N., chrysocephalus 6
UNDERWOOD, E.C., D’AMICO, J.A., ITOUA, I., pallidus 23
STRAND, H.E., MORRISON, J.C., LOUCKS, C.J., ritteri 17
ALLNUTT, T.F., RICKETTS, T.H., KURA, Y., schattatianus 22
LAMOREUx, J.F., WETTENGEL, W.W., HEDAO, P. Cereus
& KASSEM, K.R. Terrestrial Ecoregions of the anguinus 1a
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BioScience 51: 933–938. colubrinus 1a
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conservation status of cacti in the vicinity of smaragdiflorus 19
Lima, Peru. Brit. Cact. Succ. J. 14: 158–174. strausii 20
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Nazca valleys, Peru. Brit. Cact. Succ. J. 16: angosturensis 3
127–136. anguinus 1a
OSTOLAZA, C., CERONI, A., CALDERóN, N. & NOVOA, areolatus 14
S. (2003). Cacti of the Huara river valley, Peru areolatus var. herzogianus 14
a conservation study. Brit. Cact. Succ. J. 21: aureispinus 24
87–97. ayopayanus 3
RITTER, F. (1980). Kakteen in Südamerika. Band 2 azerensis 19
& 4. Self published. Spangenberg. baumannii 1a
SCHLUMPBERGER, B. & RENNER, S. (2012). baumannii ssp. anguinus 1a
Molecular phylogenetics of Echinopsis baumannii ssp. chacoanus 1a
(Cactaceae): Polyphyly at all levels and baumannii var. colubrinus 1a
convergent evolution of pollination modes and baumannii ssp. croceiflorus 1a
growth forms. Am. J. Bot. 99: 1335–1349. baumannii var. flavispinus 1a
SCHUMANN, K. (1903). Gesamtbeschreibung der baumannii var. grossei 1a
Kakteen. Neumann. Neudamm. baumannii ssp. horstii 1b
WINTER, H. (1956). Cactus seed catalogue. baumannii var. paraguariensis 1a
Frankfurt. baumannii ssp. santacruzensis 1a
brevispinus 15
brookeae 2
Index to specific epithets brookeae var. flavispinus 2
Numbers and letters refer to the numbered brookeae ssp. vulpis-cauda 2
sequence of taxa in the synopsis. Accepted names bruneispinus 1a
are indicated in bold face. buchtienii 3
buchtienii var. flavispinus 3
Bolivicereus candelilla 4
brevicaulis 18a candelilla ssp. piraymirensis 4
croceus 18a candelilla var. pojoensis 4
rufus 18a capadalensis 5
samaipatanus 18a chacoanus 1a
samaipatanus ssp. divi-miseratus 18b chacoanus var. santacruzensis 1a
samaipatanus var. divi-miseratus 18b chrysocephalus 6
samaipatanus var. multiflorus 18a clavicaulis 12
Borzicactus colademononis 24b
aureispinus 24a colubrinus 1a
aureispinus f. cristatus 24a colubrinus var. flavispinus 1a

colubrinus var. grossei 1a strausii 20
colubrinus var. smaragdiflorus 19 strausii var. fricii 9
crassicaulis 12 strausii var. jujuyensis 9
crassicaulis var. paucispinus 12 sucrensis 3
croceiflorus 1a tarijensis 9
dependens 7 tominensis 21
ferrarii 19 tominensis ssp. micropetalus 12
flavispinus 1a tupizensis 3
fusiflorus 14 tupizensis var. sucrensis 3
glaucus 11 vallegrandensis 4
glaucus var. plurispinus 11 variispinus 22
granjaensis 11 villaazuensis 13
grossei 1a villamontesii 19
herzogianus 14 villamontesii var. longiflorior 19
hildegardiae 8 viridialabastri 12
horstii 1b viridiflorus 23
hyalacanthus 9 vulpis-cauda 2
hyalacanthus ssp. tarijensis 9 wendlandiorum 2
ianthinus 14 winteri 24a
jugatiflorus 1a winteri ssp. colademono 24b
jujuyensis 9 Demnosa
jujuyensis var. fulvus 9 strausii 20
laniceps 10 Denmoza
laniceps var. plurispinus 10 strausii 20
luribayensis 11 strausii var. luteispina 9
mendozae 21 Echinopsis
micropetalus 12 baumannii 1a
morawetzianus 13 candelilla 4
morawetzianus var. pycnacanthus 13 horstii 1b
muyurinensis 4 nothohyalacantha 9
orthogonus 3 nothostrausii 20
palhuayensis 23 parviflora 14
palhuayensis var. camachoensis 23 samaipatana 18
paraguariensis 1a santacruzensis 1a
parapetiensis 19 strausii 20
parviflorus 14 tominensis 21
parviflorus var. aiquilensis 14 Hildewintera
parviflorus var. comarapanus 14 aureispina 24a
parviflorus var. herzogianus 14 colademononis 24b
piraymirensis 4 Loxanthocereus
pojoensis 4 aureispinus 24a
pungens 15 Pilocereus
pycnacanthus 13 strausii 20
reae 16 strausii f. cristatus 20
ressinianus 3 strausii var. fricii 9
ritteri 17 Seticleistocactus
rojoi 19 dependens 7
samaipatanus 18a piraymirensis 4
samaipatanus ssp. divi-miseratus 18b Winteria
santacruzensis 1a aureispina 24a
smaragdiflorus 19 Winterocereus
smaragdiflorus var. flavispinus 19 aureispinus 24a
smaragdiflorus var. gracilior 19 colademononis 24b
smaragdiflorus f. rojoi 19

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A synopsis of the genus Cleistocactus Lemaire (Cactaceae)

Article in Bradleya · November 2016

The user has requested enhancement of the downloaded ﬁle.

A synopsis of the genus Cleistocactus Lemaire (Cactaceae)

148 Bradleya 34/2016

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Habitat & Distribution: Steep cliffs and rocky

Although not rediscovered in habitat for many

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Figure 12. Cleistocactus capadalensis LM0768.01

Figure 10. Cleistocactus candelilla LM0736.01

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Figure 16. Cleistocactus dependens LM0668.01

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Etymology: from the Latin dependens; referring to

Habitat & Distribution: Hanging from cliffs in the

Although the NCL originally considered it a

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Etymology: referring to the place of its first

Habitat & Distribution: Northern dry inter

Included here are three taxa described by

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Figure 28. Cleistocactus aff. parviflorus

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Habitat & Distribution: Seasonally dry inter

Only this and one other species, C.

16. Cleistocactus reae Cárd., Cactus (Paris) 12:

A many stemmed shrub branching from the

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Cárdenas' original description of the plant as a

19. Cleistocactus smaragdiflorus (F.A.C.

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Long considerd to be an Argentinian endemic

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Figure 44. Cleistocactus winteri ssp. winteri

Figure 47. Cleistocactus winteri subsp. colade-

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Nomenclatural issues aside, C. winteri is a The discovery of this plant in 2000–2003

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Figure 48. Cleistocactus sp. An undescribed

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182 Bradleya 34/2016