Taxonomy_of_the_Cactaceae

Taxonomy of the Cactaceae

Taxonomy of the Cactaceae

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In 1984, the International Organization for Succulent Plant Study set up a working party, now called the International Cactaceae Systematics Group, to produce a consensus classification of the cactus family, down to the level of genus. Their classification has been used as the basis for systems published since the mid-1990s. Treatments in the 21st century have generally divided the family into around 125–130 genera and 1,400–1,500 species, which are then arranged in a number of tribes and subfamilies. However, subsequent molecular phylogenetic studies have shown that a very high proportion of the higher taxa (genera, tribes and subfamilies) are not monophyletic, i.e. they do not contain all of the descendants of a common ancestor. As of August 2023, the internal classification of the family Cactaceae remained uncertain and subject to change. A classification incorporating many of the insights from the molecular studies was produced by Nyffeler and Eggli in 2010.[1]

A collection of cultivated cacti

Overview

The classification of the family Cactaceae remains uncertain as of August 2023. Since the mid-1990s, the system produced by the International Cactaceae Systematics Group (ICSG) of the International Organization for Succulent Plant Study has been used as the basis of many published classifications. Detailed treatments produced in the 21st century have divided the family into around 125–130 genera and 1,400–1,500 species, which are then arranged into a number of tribes and subfamilies.[2][3][4]

The ICSG classification of the family recognizes four subfamilies: Pereskioideae (consisting only of the genus Pereskia), Opuntioideae, Maihuenioideae (consisting only of the genus Maihuenia) and Cactoideae.[2] Molecular phylogenetic studies suggest that Pereskia is not monophyletic (i.e. its species are not the complete set of descendants of a common ancestor), so that Pereskioideae is not monophyletic although the three other subfamilies are.[4][5] The Bayesian consensus cladogram from a 2005 study is shown below:[5]

Cactaceae

Five tribes have been recognized within the subfamily Opuntioideae: Tephrocacteae, Pterocacteae, Austrocylindropuntieae, Cylindropuntieae and Opuntieae.[6] All but the first, Tephrocacteae, were shown to be "essentially monophyletic" in molecular phylogenetic study in 2009. A summary of the consensus Bayesian tree is shown below (tribes are bold; the number of species in the study is shown in parentheses).[7]

Puna (1), Maihueniopsis (3), Cumulopuntia (1) = Tephrocacteae (part)

Pterocactus (6) = Pterocacteae

terete‑stemmed opuntioids
flat‑stemmed opuntioids

Many of the genera within the Opuntioideae are not monophyletic. Maihueniopsis is highly polyphyletic, appearing in four separate lineages. The two largest genera within the subfamily, Opuntia and Cylindropuntia are also not monophyletic. The classification of the Opuntioideae is thus uncertain as of March 2012; Griffith and Porter say that changes in classification will require "broad information (of multiple data types) regarding all species of opuntioid cacti".[8]

The ICSG classification divides the subfamily Cactoideae into nine tribes. However, phylogenetic research has not supported most of these tribes, nor even the genera of which they are composed. A 2011 study found that "an extraordinarily high proportion of genera" were not monophyletic, including 22 (61%) of the 36 genera in the subfamily Cactoideae sampled in the research.[4] Of the nine tribes recognized within Cactoideae, one, Calymmantheae, comprises a single genus, Calymmanthium.[2] Of the remaining eight, only two (Cacteae and Rhipsalideae) have been shown to be monophyletic. A summary of the cladograms for the Cactoideae presented in a 2011 paper is shown below (ICSG tribes in bold).[9]

Cactoideae

Blossfeldia

Cacteae

core Cactoideae

Calymmantheae (i.e. Calymmanthium), species of Copiapoa and Frailea

core Cactoideae I

including members of Pachycereeae, Hylocereeae and Browningieae

core Cactoideae II

Rhipsalideae

other core Cactoideae II, including members of Notocacteae, Trichocereeae, Cereeae and Browningieae

The classification of the Cactaceae thus remains subject to change, from the genus level upwards. In 2016, David Hunt wrote that "the advent of molecular systematic studies in the past two decades will hopefully have a stabilizing effect on classification in due course, but has so far had the opposite effect".[10]

Nyffler and Eggli (2010) classification

In 2010, Nyffler and Eggli produced a classification based on an explicit phylogeny. The broad outlines of their cladogram agree with that of Hern et al. (2011), shown above, although some details are different. Their classification corresponds to the cladogram shown below.[1]

Cactaceae
Pereskioideae
Maihuenioideae
Opuntioideae
Cactoideae

The table below shows how the genera recognized by Nyffler and Eggli are placed into their subfamilies and tribes (they also use some subtribes, not shown here). The column headed "Older tribe (if different)" shows the ICSG classification of the subfamily Cactoideae (as presented by Anderson in 2001[11]) with a 2002 classification of the subfamily Opuntioideae.[6][12] There are differences in the genera recognized in the systems; where the older system does not recognize the genus, "–" is shown.

More information Systematic order, Genus ...

Earlier classification

Unless otherwise indicated, the subfamily placement of the genera listed here is based on Anderson's 2001 presentation of the ICSG classification, as is the tribal placement of the genera of the subfamily Cactoideae.[11] The division of the subfamily Opuntioideae into tribes is additional to the ICSG system.[6][12]

Subfamily Pereskioideae

Pereskia aculeata
synonyms: Peirescia Zucc. (orth. var.), Peireskia Steud. (orth. var.), Perescia Lem. (orth. var.), Rhodocactus (A.Berger) F.M.Knuth

Subfamily Opuntioideae

Tribe Austrocylindropuntieae

Tribe Cylindropuntieae

including Corynopuntia F.M.Knuth, Micropuntia Daston
synonyms: Peireskiopsis Vaupel (orth. var.)

Tribe Opuntieae

Opuntia ficus-indica
synonyms: Airampoa Fric, Cactodendron Bigelow, nom. inval., Cactus Lem., Chaffeyopuntia Fric & Schelle, Clavarioidia Kreuz. (nom. inval.), Ficindica St.-Lag., Nopalea Salm-Dyck, Parviopuntia Soulaire & Marn.-Lap. (nom. inval.), Phyllarthus Neck. ex M.Gómez (nom. inval.), Salmiopuntia Fric (nom. inval.), Subulatopuntia Fric & Schelle, Tunas Lunell, Weberiopuntia Fric
including Nopalea Salm-Dyck

Tribe Pterocacteae

Tribe Tephrocacteae

including Puna R.Kiesling
synonyms: Pseudotephrocactus Fric

Subfamily Maihuenioideae

Maihuenia poeppigii

Subfamily Cactoideae

Tribe Browningieae

synonyms: Neocardenasia Backeb.

Tribe Cacteae

synonyms: Anhalonium Lem., Neogomesia Castañeda, Neogomezia Buxb. (orth. var.), Roseocactus A.Berger, Stromatocactus Karw. ex Rümpler (nom. inval.)
Astrophytum myriostigma
synonyms: Astrophyton Lawr. (orth. var.), Maierocactus E.C.Rost
synonyms: Aulacothele Monv. (nom. inval.), Cumarinia Buxb., Glandulifera (Salm-Dyck) Fric, Lepidocoryphantha Backeb., Roseia Fric (nom. inval.)
synonyms: Brittonrosea Speg., Echinofossulocactus Lawr., Homalocephala Britton & Rose
synonyms: Cochiseia W.H.Earle, Escobesseya Hester, Fobea Fric (nom. inval.), Neobesseya Britton & Rose
synonyms: Bisnaga Orcutt, Brittonia C.A.Armstr. (nom. inval.)
Mammillaria elongata
  • Mammillaria Haw. – fishhook cactus, globe cactus, pincushion cactus, bird's-nest cactus
synonyms: Bartschella Britton & Rose, Cactus L., Chilita Orcutt, Cochemiea (K.Brandegee) Walton, Dolichothele (K.Schum.) Britton & Rose, Ebnerella Buxb., Haagea Fric, Krainzia Backeb., Lactomammillaria Fric (nom. inval.), Leptocladia Buxb., Leptocladodia Buxb., Mamillaria F.Rchb. (orth. var.), Mamillopsis (E.Morren) F.A.C.Weber ex Britton & Rose, Mammariella Shafer (nom. inval.), Mammilaria Torr. & A.Gray (orth. var.), Neomammillaria Britton & Rose, Oehmea Buxb., Phellosperma Britton & Rose, Porfiria Boed., Pseudomammillaria Buxb., Solisia Britton & Rose
synonyms: Napina Fric (nom. inval.), Pseudosolisia Y.Itô (nom. inval.)
synonyms: Navajoa Croizat, Pilocanthus B.W.Benson & Backeb.
synonyms: Encephalocarpus A.Berger
Sclerocactus glaucus
synonyms: Ancistrocactus Britton & Rose, Coloradoa Boissev. & C.Davidson, Glandulicactus Backeb., Toumeya Britton & Rose
synonyms: Echinofossulocactus Britton & Rose, Efossus Orcutt (orth. var.)
synonyms: Hamatocactus Britton & Rose, Thelomastus Fric (nom. inval.)
synonyms: Gymnocactus Backeb., Normanbokea Kladiwa & Buxb., Rapicactus Buxb. & Oehme

Tribe Calymmantheae

synonyms: Diploperianthium F.Ritter (nom. inval.)

Tribe Cereeae

synonyms: Digitorebutia Frič & Kreuz. ex Buining, Mediolobivia Backeb.
synonyms: Pierrebraunia Esteves
synonyms: Mirabella F.Ritter, Piptanthocereus (A.Berger) Riccob., Subpilocereus Backeb.
synonyms: Floribunda F.Ritter
synonyms: Buiningia Buxb.
Melocactus matanzanus
synonyms: Cactus Britton & Rose
synonyms: Austrocephalocereus Backeb., Siccobaccatus P.J.Braun & Esteves
synonyms: Pilocereus K.Schum., Pseudopilocereus Buxb.

Tribe Hylocereeae

synonyms: Aporocactus Lem., Aporocereus Fric & Kreuz. (orth. var.), Bonifazia Standl. & Steyerm., Chiapasia Britton & Rose, Disisocactus Kunze (orth. var.), Disocereus Fric & Kreuz. (orth. var.), Heliocereus (A.Berger) Britton & Rose, Lobeira Alexander, Mediocereus Fric & Kreuz. (orth. var.), Nopalxochia Britton & Rose, Pseudonopalxochia Backeb., Trochilocactus Linding., Wittia K.Schum., Wittiocactus Rauschert
Epiphyllum anguliger
synonyms: Phyllocactus Link, Phyllocereus Miq.
  • Hylocereus (A.Berger) Britton & Rose – nightblooming cactus
synonyms: Wilmattea Britton & Rose
synonyms: Cryptocereus Alexander, Deamia Britton & Rose, Marniera Backeb., Mediocactus Britton & Rose, Strophocactus Britton & Rose, Strophocereus Fric & Kreuz. (orth. var.)
synonyms: Eccremocactus Britton & Rose, Eccremocereus Fric & Kreuz. (orth. var.), Werckleocereus Britton & Rose

Tribe Notocacteae

synonyms: Pilocopiapoa F.Ritter
Eriosyce esmeraldana
synonyms: Ceratistes Labour. (nom. inval.), Chileniopsis Backeb., Chileocactus Fric (nom. inval.), Chileorebutia Fric (nom. inval.), Chiliorebutia Fric (orth. var.), Delaetia Backeb., Dracocactus Y.Itô (nom. inval.), Euporteria Kreuz. & Buining, Hildmannia Kreuz. & Buining, Horridocactus Backeb., Islaya Backeb., Neochilenia Backeb. ex Dölz, Neoporteria Britton & Rose, Neoporteria Backeb., Neotanahashia Y.Itô, Nichelia Bullock (nom. inval.), Pyrrhocactus (A.Berger) A.W.Hill, Rodentiophila F.Ritter ex Backeb., Thelocephala Y.Itô
synonyms: Philippicereus Backeb.
synonyms: Acanthocephala Backeb., Brasilicactus Backeb., Brasiliparodia F.Ritter, Brasilocactus Fric (nom. inval.), Chrysocactus Y.Itô (nom. inval.), Dactylanthocactus Y.Itô, Eriocactus Backeb., Eriocephala Backeb., Friesia Fric (nom. inval.), Hickenia Britton & Rose, Malacocarpus Salm-Dyck, Microspermia Fric, Neohickenia Fric, Notocactus (K.Schum.) Fric, Sericocactus Y.Itô, Wigginsia D.M.Porter

Tribe Pachycereeae

synonyms: Dendrocereus Britton & Rose, Monvillea Britton & Rose
synonyms: Bergerocereus Fric & Kreuz. (orth. var.)
Carnegiea gigantea
synonyms: Haseltonia Backeb., Neodawsonia Backeb., Pilocereus Lem.
synonym: Corryocereus Fric & Kreuz. (orth. var.), Erdisia Britton & Rose, Eulychnocactus Backeb. (nom. inval.)
synonyms: Morangaya G.D.Rowley, Wilcoxia Britton & Rose
synonyms: Neoabbottia Britton & Rose
[= Myrtillocactus × Bergerocactus]
synonyms: Myrtillocereus Fric & Kreuz. (orth. var.)
Pachycereus pringlei
synonyms: Pseudomitrocereus Bravo & Buxb., Rooksbya Backeb.
[= Pachycereus × Bergerocactus]
synonyms: Backebergia Bravo, Lemaireocereus Britton & Rose, Lophocereus (A.Berger) Britton & Rose, Marginatocereus (Backeb.) Backeb., Mitrocereus (Backeb.) Backeb., Pterocereus T.MacDoug. & Miranda
synonyms: Cullmannia Distefano, Neoevansia W.T.Marshall, Nyctocereus (A.Berger) Britton & Rose
synonyms: Chichipia Backeb. (nom. inval.), Heliabravoa Backeb.
synonyms: Hertrichocereus Backeb., Isolatocereus Backeb., Isolatocereus (Backeb.) Backeb., Machaerocereus Britton & Rose, Marshallocereus Backeb., Neolemaireocereus Backeb., Rathbunia Britton & Rose, Ritterocereus Backeb.

Tribe Rhipsalideae

Hatiora salicornioides
synonyms: Epiphyllopsis Backeb. & F.M.Knuth, Hariota DC., Pseudozygocactus Backeb., Rhipsalidopsis Britton & Rose
synonyms: Acanthorhipsalis (K.Schum.) Britton & Rose, Acanthorhipsalis Kimnach, Pfeiffera Salm-Dyck
synonyms: Cassytha Mill., Erythrorhipsalis A.Berger, Hariota Adans., Lymanbensonia Kimnach
synonyms: Epiphyllanthus A.Berger, Epiphyllum Pfeiff., Opuntiopsis Knebel (nom. inval.), Zygocactus K.Schum., Zygocereus Fric & Kreuz. (orth. var.)

Tribe Trichocereeae

synonyms: Spinicalycium Fric (nom. inval.)
Cleistocactus
synonyms: Akersia Buining, Bolivicereus Cárdenas, Borzicactella H.Johnson ex F.Ritter, Borzicactus Riccob, Borzicereus Fric & Kreuz. (orth. var.), Cephalocleistocactus F.Ritter, Cleistocereus Fric & Kreuz. (orth. var.), Clistanthocereus Backeb., Demnosa Fric, Gymnanthocereus Backeb., Hildewintera F.Ritter, Loxanthocereus Backeb., Maritimocereus Akers, Pseudoechinocereus Buining (nom. inval.), Seticereus Backeb., Seticleistocactus Backeb., Winteria F.Ritter, Winterocereus Backeb.
synonyms: Acantholobivia Backeb., Acanthopetalus Y.Itô, Andenea Fric (nom. inval.), Aureilobivia Fric, nom. inval., Chamaecereus Britton & Rose, Chamaelobivia Y. Itô (nom. inval.), Cinnabarinea Fric ex F.Ritter, Echinolobivia Y.Itô (nom. inval.), Echinonyctanthus Lem., Furiolobivia Y.Itô (nom. inval.), Helianthocereus Backeb, Heterolobivia Y.Itô (nom. inval.), Hymenorebulobivia Fric (nom. inval.), Hymenorebutia Fric ex Buining, Leucostele Backeb., Lobirebutia Fric (nom. inval.), Lobivia Britton & Rose, Lobiviopsis Fric (nom. inval.), Megalobivia Y.Itô (nom. inval.), Mesechinopsis Y.Itô, Neolobivia Y.Itô, Pilopsis Y.Itô (nom. inval.), Pseudolobivia (Backeb.) Backeb., Rebulobivia Fric (nom. inval.), Salpingolobivia Y.Itô, Scoparebutia Fric & Kreuz. ex Buining, Setiechinopsis (Backeb.) de Haas, Soehrensia Backeb., Trichocereus (A.Berger) Riccob.
synonyms: Binghamia Britton & Rose, Pseudoespostoa Backeb., Thrixanthocereus Backeb.,
synonyms: Gerocephalus F.Ritter
synonyms: Zehntnerella Britton & Rose
Gymnocalycium moserianum
synonyms: Brachycalycium Backeb.
synonyms: Floresia Krainz & F.Ritter ex Backeb. (nom. inval.), Haageocactus Backeb. (nom. inval.), Lasiocereus F.Ritter, Neobinghamia Backeb., Peruvocereus Akers
[= Haageocereus × Espostoa]
synonyms: Eriocereus (A.Berger) Riccob., Roseocereus Backeb.
synonyms: Eomatucana F.Ritter
synonyms: Arequipa Britton & Rose, Arequipiopsis Kreuz. & Buining, Morawetzia Backeb., Submatucana Backeb.
Rebutia flavistyla
synonyms: Aylostera Speg., Bridgesia Backeb., Cylindrorebutia Fric & Kreuz., Digitorebutia Fric & Kreuz., Echinorebutia Fric (nom. inval.), Eurebutia Fric (nom. inval.), Gymnantha Y.Itô, Mediolobivia Backeb., Mediorebutia Fric (nom. inval.), Neogymnantha Y.Itô, Reicheocactus Backeb., Setirebutia Fric & Kreuz. (nom. inval.), Spegazzinia Backeb., Sulcorebutia Backeb., Weingartia Werderm.
synonyms: Meyenia Backeb.

Notes and references

  1. [1]
    Nyffeler, R. & Eggli, U. (2010), "A farewell to dated ideas and concepts: molecular phylogenetics and a revised suprageneric classification of the family Cactaceae", Schumannia, 6: 109–149, doi:10.5167/uzh-43285, S2CID 89650660
  2. [2]
    Anderson 2001, pp. 99–103
  3. [3]
    Hunt, D.R., ed. (2006), The New Cactus Lexicon (two volumes), Milborne Port: dh books, ISBN 978-0-9538134-4-5, cited in Bárcenas, Yesson & Hawkins 2011
  4. [4]
    Bárcenas, Rolando T.; Yesson, Chris & Hawkins, Julie A. (2011), "Molecular systematics of the Cactaceae", Cladistics, 27 (5): 470–489, doi:10.1111/j.1096-0031.2011.00350.x, PMID 34875796, S2CID 83525136
  5. [5]
    Edwards, Erika J.; Nyffeler, Reto & Donoghue, Michael J. (2005), "Basal cactus phylogeny: implications of Pereskia (Cactaceae) paraphyly for the transition to the cactus life form", American Journal of Botany, 92 (7): 1177–1188, doi:10.3732/ajb.92.7.1177, PMID 21646140
  6. [6]
    Wallace, R.S. & Dickie, S.L. (2002), "Systematic implication of chloroplast DNA sequence variation in subfam. Opuntioideae (Cactaceae)", Succulent Plant Research, 6: 9–24, cited in Griffith & Porter 2009
  7. [7]
    Griffith, M. Patrick & Porter, J. Mark (2009), "Phylogeny of Opuntioideae (Cactaceae)", International Journal of Plant Sciences, 170 (1): 107–116, doi:10.1086/593048, S2CID 85142301. Tree based on fig. 1 with tribes from fig. 2.
  8. [8]
    Griffith & Porter 2009, p. 114
  9. [9]
    Hernández-Hernández, Tania; Hernández, Héctor M.; De-Nova, J. Arturo; Puente, Raul; Eguiarte, Luis E. & Magallón, Susana (2011), "Phylogenetic relationships and evolution of growth form in Cactaceae (Caryophyllales, Eudicotyledoneae)", American Journal of Botany, 98 (1): 44–61, doi:10.3732/ajb.1000129, PMID 21613084
  10. [10]
    Hunt, David (2016), CITES Cactaceae Checklist (PDF) (3rd ed.), Royal Botanic Gardens, ISBN 978-0-9933113-2-1, retrieved 2017-03-30
  11. [11]
    Anderson 2001, pp. 100–103
  12. [12]
    "Family: Cactaceae Juss., nom. cons. subfam. Opuntioideae", Germplasm Resources Information Network, United States Department of Agriculture, 2004-02-13, archived from the original on 2012-08-25, retrieved 2012-03-26
  13. [13]
    Bárcenas, Rolando, Héctor M. Hernández, Patricia Hernández-Ledesma, and Luis Manuel Montoya Gómez (2021). Chichimecactus (Cactoideae, Cactaceae), a new genus based on molecular characterisation of highly endangered Strombocactus species. Phytotaxa Vol. 512 No. 3: 27 July 2021. DOI: https://doi.org/10.11646/phytotaxa.512.3.2
  14. [14]
    Kiesling, R. & Ferrari, O. (2003), "Yavia cryptocarpa – conservation action on a new and interesting cactus", British Cactus and Succulent Journal, 21 (1): 20–25

Bibliography
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