1

The CITES Cactaceae Checklist

CITES: The Convention on International Trade in Endangered Species
Signatories: 180 Countries
[1] Apart from Opuntia in the narrow sense and various cultivars, the entire cactus family
is subject to the provisions of CITES Appendix II. That basically means that permits are
always needed to export or import cacti.
Various genera and species are covered by the more stringent Appendix I, but in South
America only Discocactus, some Melocactus species and Uebelmannia in Brazil.
[2] We all need stable scientific names for use in agriculture, conservation, forestry,
horticulture, medical research etc., especially at the level of genus, because thats how
animal and plant names are indexed.
Stable names are very important to CITES because name changes in their Checklists may
lead to costly updates of their permit systems and confusion for enforcement agencies
and traders, importers and users generally.
Unfortunately, the generic classification of cacti has been notoriously unstable since
Britton & Rose inflated the modest number accepted by Schumann by over 100.
[3] Bergers system was more like Schumanns but was soon submerged by the flood of
publications by Backeberg. His phenomenal output is said to have included new species
seen from a moving train.
[4] By the time of his death in 1966, Backeberg had added another 100+ genera. He also
described an enormous number of species without ever preserving a nomenclatural type.
Franz Buxbaum made valuable surveys of morphological features, among which seed
morphology has proved particularly useful in cactus taxonomy, especially since the
advent of the electron microscope.
After Backeberg's death a more conservative approach began to prevail and by 1993 the
total recognized by Barthlott & Hunt for Kubitskis Families and Genera of Flowering
Plants had shrunk to 98.
[5] A working party to seek a consensus on the generic classification of the family was set
up in 1984 under the auspices of the International Organization for Succulent Plant
Study. A few years later it was tasked by CITES to produce a Standard Reference for the
use of those implementing the CITES Convention. The first edition of the Checklist was
published in 1992 and the second in 1999.
[6] In 2000, the working party was re-named the International Cactaceae Systematics
Group. Having worked together at more than 20 discussion meetings, we published the
New Cactus Lexicon, with illustrations of over 90 per cent. of the species we could accept.
We accepted the same number of genera as Britton & Rose, but not all the same ones!
David Hunt 2015 [Peru talk]

This year Joel Lod has published what he calls the new classification of Cacti mainly
based on molecular data and explained. He accepts many more genera than my group
but has not done any of the research and does not understand that molecular evidence
does not dictate that every small monophyletic group should be treated as a genus.
To achieve nomenclatural stability we need to make the most of the inherent flexibility of
the traditional hierarchical system, like Schumann and Berger. By using the traditional
infrageneric categories of subgenus etc, changes of classification can be proposed and
differences of opinion can be held without creating more names and synonyms.
[7] The Checklist Compilers task in the age of Molecular Systematics: To reconcile
phylogenetic hypotheses with practical taxonomy and nomenclatural stability.
These days genera are supposed to be monophyletic, thats to say all the species should
be directly related by ancestry. Lets consider three examples affecting the cacti of Peru:
First, the case of Opuntia, the largest genus in Britton & Rose's monograph:
In 1986, The majority view of the Working Party was that there was not yet sufficient
evidence for a stable infrageneric classification of Opuntia or for the recognition of
potential segregates such as Cylindropuntia and Tephrocactus. (Hunt & Taylor eds.
1986). This treatment was followed in the first two editions of the CITES Checklist, but by
2002 molecular and seed micromorphological evidence was sufficient to persuade a
majority of the Working Party to accept a division of the genus into 10 genera and five
Tribes, since reduced to three that broadly correspond to the three subgenera into which
Britton & Rose divided Opuntia.
[8] What would you do if you were the compiler of the new Checklist?
The dissection of Opuntia has certainly encouraged further research on the subfamily.
But, from the point of view of nomenclatural stability, it seems to have served as a
precedent for dividing other genera, large and small, rather than treating the component
infrageneric groups as subgenera etc.
For the purpose of the CITES Checklist, at least, the door still remains open to treat
Britton & Rose's three subgenera as genera and the rest of the segregates as subgenera.
Relatively few new combinations would be needed.
[9] Taking the tribe Tephrocacteae first, a recent molecular study (Ritz et al. 2012) found
several individual species to be misclassified, one genus paraphyletic and another
polyphyletic, which means of mixed ancestry . [See Table at end].
[10] All the little genera in tribe Tephrocacteae could be accommodated as subgenera
difficulty in the genus Tephrocactus itself (subg. Tephrocactus Britton & Rose).
[11] The much-misunderstood Austrocylindropuntia lagopus proved to be basal to the
Austrocylindropuntieae group, making that genus paraphyletic unless Cumulopuntia was
included as well, or else A. lagopus was given a separate genus. After much discussion, at
David Hunt 2015 [Peru talk]

a meeting with the principal researcher, Dr Christiane Ritz, we opted for the latter option,
at least for the time being.
[12] The genus Puna was proposed for 3 geophytic species. Each of them has since been
shown by seed morphology and molecular data to belong a different genus.
[13] Second example: The case for Lymanbensonia Kimnach: In the second edition of the
CITES Checklist, these two Peruvian species were included in Lepismium, following
Barthlott & Taylor (1995/Brdl. 13: 4448). But molecular evidence (Nyffeler 2000)
showed that Lepismium and Pfeiffera are not closely related, Pfeiffera being closer to
Corryocactus etc than to the Rhipsalis group. So they were transferred to Pfeiffera.
Then a broader molecular study by a student of Professor Barthlott, Nadja Korotkova
(2010), showed that they and two Bolivian species of Pfeiffera form a separate clade,
closer to Calymmanthium, and differ from Pfeiffera in their flowers, which have a distinct
tube and spineless fruits. Nadja proposed reinstating the genus Lymanbensonia.
Rhipsalis riocampanensis Madsen et al. (2004) is thought to be the same as
Lymanbensonia brevispina, extending the range of the genus to Ecuador.
[14] Third example: Tribe Trichocereeae: Molecular evidence (Schlumpberger & Renner
2012) suggests that Echinopsis would only be monophyletic if it was expanded to include
most of the other hitherto recognized genera of Trichocereeae. This radical option is
favoured by some authors (Anceschi & Magli 2013; Molinari-Novoa & Mayta 2015) but is
not very practical and would require changing many names.
The alternative, proposed by Schlumpberger, would be to reinstate Lobivia, Trichocereus
and several mostly small genera currently subsumed in Echinopsis.
Actually, Echinopsis in the narrow sense doesnt occur in Peru. All the Peruvian species
listed in the 1999 edition of the CITES Checklist have alternative names in Lobivia or
Trichocereus, so I would recommend using them.
The other native genera form two distinct clades in the Schlumpberger system. This
accords with the hypothesis that adaptations to bird-pollination evolved independently
in the two main lineages. Consequently the inclusion of Borzicactus in Cleistocactus,
adopted in earlier editions of the CITES Checklist, has to be rejected.
[15] The Cleistocactus and Borzicactus clades: As in the Opuntioideae, opinions may
differ on how many genera to accept: 1. Recognize all 16 genera; 2. Recognize one genus
only per clade; 3. Select two or more genera in each clade:
Cleistocactus clade: Cleistocactus, Samaipaticereus*. Vatricania*, Weberbauerocereus,
Yungasocereus*
Borzicactus clade: Borzicactus, Espostoa, Haageocereus, Lasiocereus, Loxanthocereus,
Matucana, Mila, Oreocereus, Oroya, Pygmaeocereus, Rauhocereus* [* = monotypic]

David Hunt 2015 [Peru talk]

Kimnach (1960) proposed expanding Borzicactus to include Arequipa, Loxanthocereus

Matucana and Oreocereus. Dr Schlumpberger is doing further research on the
Borzicactus clade. His initial results suggest that Loxanthocereus is closer to
Haageocereus than to Borzicactus.
[16] Ritter (1981) reported numerous suspected indigenous intergeneric hybrids in the
Peruvian Trichocereae, some of them apparently intercladal: Cleistocana (Cleistocactus
Matucana), Espocana (Espostoa Matucana) , Espostingia (Espostoa Rauhocereus),
Espostocactus (Espostoa Cleistocactus), Haagespostoa (Haageocereus Espostoa),
Maturoya (Matucana Oroya), Oreocana (Oreocereus Matucana), Weberbostoa
(Weberbauerocereus Espostoa), Yungastocactus (Yungasocereus Cleistocactus).
Basic evidence from cpDNA is needed but would only reflect the maternal parentage, as
would seed micromorphology. Cytogenetic evidence (ploidy etc) also needed. If proven,
such hybrids could be taken as an indication of the close relationships of the putative
genera in the tribe, and an argument in favour of recombining them.
[17] Besides those in Trichocereeae there are actually quite a lot of cacti that could prove
to be intergeneric hybrids. Echinocactus grusonii is probably the most famous.
[18] Back to Peru and congratulations to Carlos Ostolaza on his latest book. We are very
largely in agreement on which genera to accept. Currently we include Loxanthocereus in
Borzicactus but as I mentioned it may be closer to Haageocereus. The problem with
Monvillea is that the type species was an Acanthocereus from Colombia. So we consider
that Monvillea is a misapplied name and Praecereus the correct name. In any event
molecular evidence now suggests Praecereus could or should be included in Cereus.
As I mentioned earlier, the name Lymanbensonia has recently been reinstated for the
two Peruvian species of Pfeiffera and Punotia is an alternative name for Austrocylindropuntia lagopus. Molecular evidence excludes Strophocactus wittii from
Selenicereus. Tephrocactus verschaffeltii occurs in the Machu Picchu valley.*
[19] Topics such as these are raised for information and discussion in the normally twiceyearly Bulletin that I started in 1996 and circulate to those helping with the compilation
of the CITES Checklist and the New Cactus Lexicon and also to others interested. Initially
called 'Cactaceae Consensus Initiatives', we changed its name to Cactaceae Systematics
Initiatives' in 2000. [Who knows the species on the cover on no. 17?]
[20] The current issue (no. 33) also touches on the limitations of chloroplast DNA
evidence and the role of hybridization in cactus evolution. These are topics that deserve
more attention from molecular systematists, with the help of cytogeneticists.
[21] That reminds me to say that if members of SPECS who are involved in research
projects concerning succulent plants or would like to start one but are, as we say
strapped for cash, the IOS, of which Im Honorary Secretary, may be able to help. Just
contact me with an idea of what you want to do. Thank you for your attention!
David Hunt 2015 [Peru talk]

*Hoxey, P. (2014). Tephrocactus verschaffeltii - A surprising addition to the cactus flora of

Peru. Bradleya 32: 92-98.

[Table for no. 9]

Tribe Tephrocacteae (Ritz et al. 2012)
Austrocylindropuntia Backeb.
lagopus

= Punotia

verschaffeltii = Tephrocactus
Cumulopuntia Ritter 1980
recurvata

= Tephrocactus

Maihueniopsis Speg. 1925

nigrispinus = Tephrocactus
subterranea = Cumulopuntia
Puna Kiesling 1982
clavarioides = Maihueniopsis
bonniae

= Tephrocactus

subterranea = Cumulopuntia

David Hunt 2015 [Peru talk]