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Echinocereeae
Echinocereus stramineus
Saguaro, Carnegiea gigantea
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Order:	Caryophyllales
Family:	Cactaceae
Subfamily:	Cactoideae
Tribe:	Echinocereeae Buxbaum
Genera
See text.

The Echinocereeae are a tribe of cacti in the subfamily Cactoideae. Since 2006, the tribe has included the former tribe Pachycereeae in many treatments of cactus classification. The exact circumscription of the tribe has been subject to considerable change, particularly since molecular phylogenetic approaches have been used in determining classifications, and remains uncertain. The tribe includes large treelike species, such as the saguaro (Carnegiea gigantea), as well as shorter shrubby species. Most members of the tribe are found in desert regions, particularly in Mexico and the southwestern United States.

Description

The tribe includes large treelike species, as well as shorter shrubby species. Some species can grow to be over 15 m (50 ft) tall, like the saguaro (Carnegiea gigantea) and Cephalocereus macrocephalus (syn. Neobuxbaumia macrocephala). Their stems are ribbed and columnar, not divided into segments. Most have flowers that open at night.

Taxonomy

In 1958, Franz Buxbaum published a division into tribes of what is now the subfamily Cactoideae, then Cereoideae. Two of these tribes were Pachycereeae (which he spelt "Pachycereae") and Echinocereeae (which he spelt "Echinocereae"). Pachycereeae was a newly published taxon; Echinocereeae was an elevation of the subtribe Echinocereinae ("Echinocereanae") established by Britton and Rose in 1922. Studies based on chloroplast DNA showed that Echinocereus, the type genus of the Echinocereeae, was nested within the Pachycereeae, as then circumscribed, and Pachycereeae was expanded to include Echinocereeae. In 2006, David Hunt and the International Cactaceae Systematics Group used the name Echinocereeae rather than Pachycereeae for the tribe. This has been followed in much subsequent work, including a major 2011 molecular phylogenetic analysis of the family Cactaceae. However, some authors continue to use the name Pachycereeae for the tribe.

Phylogeny

Genera placed at one time or another within the Echinocereeae (or Pachycereeae) fell within the "Core Cactoideae I" clade in the 2011 molecular phylogenetic study by Barcenas et al. Possible phylogenetic relationships within this clade are shown in the cladogram below, based on Guerrero et al. (2019) and revisions to the Hylocereeae in 2017 and 2020.

Genera

"Core Echinocereeae" recovered in molecular phylogenetic analyses in 2005 and 2011 contains the following genera, often only in part compared to earlier generic circumscriptions. Some genera were further placed in two subtribes, Pachycereinae and Echinocereinae. The position of some genera remained unclear, as of October 2023. Brachycereus was not included in many studies, but has been placed with Armatocereus and Jasminocereus, both included in Echinocereeae.

Image	Genus	Subtribe
	Armatocereus Backeb.	-
	Bergerocactus Britton & Rose	Pachycereinae
	Brachycereus Britton & Rose 1920	-
	Carnegiea Britton & Rose	Pachycereinae
	Cephalocereus Pfeiff.	Pachycereinae
	Echinocereus Engelm.	Echinocereinae
	Escontria Rose	–
	Jasminocereus Britton & Rose	-
	Mitrocereus (Backeb.) Backeb.	Echinocereinae
	Myrtillocactus Console	Echinocereinae
	Neoraimondia Britton & Rose
	Pachycereus (A.Berger) Britton & Rose	Pachycereinae
	Pterocereus T.MacDoug. & Miranda	Pachycereinae
	Peniocereus (A.Berger) Britton & Rose	Pachycereinae
	Polaskia Backeb.	Echinocereinae
	Stenocereus (A.Berger) Riccob. (including Isolatocereus)	Echinocereinae
	Corryocactus Britton & Rose ("outside of a well-supported larger clade")	–
	Deamia Britton & Rose	–
	Leptocereus (A.Berger) Britton & Rose	–
	Lophocereus Britton & Rose, when it is separated from Pachycereus	Pachycereinae
	Strophocactus Britton & Rose	–
	Morangaya G.D.Rowley	-

The 2011 study found the tribe Hylocereeae to be embedded within a wider Echinocereeae, but studies in 2017 and 2020 asserted the independence of the Hylocereeae, including within it two genera placed within Pachycereeae in earlier classifications: Acanthocereus and Pseudoacanthocereus (sunk into Strophocactus).

Distribution and habitat

Members of the Echinocereeae are native from the southwestern United States, through Mexico, Central America and the Caribbean to northern South America. Many are classic columnar desert cacti of Mexico and the southwestern US, like the saguaro (Carnegiea gigantea). A few have a wider distribution: Stenocereus griseus is found from northeastern Mexico to Colombia.

References

1. ^ Anderson, Edward F. (2001), The Cactus Family, Pentland, Oregon: Timber Press, ISBN 978-0-88192-498-5, p. 103
2. ^ Anderson (2001), p. 138
3. Anderson (2001), p. 477
4. Buxbaum, Franz (1958), "The Phylogenetic Division of the Subfamily Cereoideae, Cactaceae", Madroño, 14 (6): 177–206, JSTOR 41422934
5. Reveal, James L. (2011), "Indices Nominum Supragenericorum Plantarum Vascularium – E", PlantSystematics.org, retrieved 2018-03-11
6. Zimmerman, Allan D. & Parfitt, Bruce D., "Echinocereus", in Flora of North America Editorial Committee (ed.), Flora of North America (online), eFloras.org, retrieved 2021-03-16
7. Hunt, David, ed. (2006), The New Cactus Lexicon, Milborn Port: Dh Books, ISBN 978-0-9538134-5-2
8. ^ Barcenas, 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
9. ^ Copetti, Dario; Búrquez, Alberto; Bustamante, Enriquena; Charboneau, Joseph L. M.; Childs, Kevin L.; Eguiarte, Luis E.; Lee, Seunghee; Liu, Tiffany L.; McMahon, Michelle M.; Whiteman, Noah K.; Wing, Rod A.; Wojciechowski, Martin F. & Sanderson, Michael J. (2017), "Extensive gene tree discordance and hemiplasy shaped the genomes of North American columnar cacti", Proceedings of the National Academy of Sciences, 114 (45): 12003–12008, doi:10.1073/pnas.1706367114, PMC 5692538, PMID 29078296
10. Guerrero, Pablo C.; Majure, Lucas C.; Cornejo-Romero, Amelia & Hernández-Hernández, Tania (2019-01-07), "Phylogenetic Relationships and Evolutionary Trends in the Cactus Family", Journal of Heredity, 110 (1): 4–21, doi:10.1093/jhered/esy064, PMID 30476167
11. ^ Korotkova, Nadja; Borsch, Thomas & Arias, Salvador (2017). "A phylogenetic framework for the Hylocereeae (Cactaceae) and implications for the circumscription of the genera". Phytotaxa. 327 (1): 1–46. doi:10.11646/phytotaxa.327.1.1.
12. ^ Martínez-Quezada, Daniel M.; Arias, Salvador; Korotkova, Nadja & Terrazas, Teresa (2020), "The phylogenetic significance of the stem morpho-anatomy in the Hylocereeae (Cactoideae, Cactaceae)", Plant Systematics and Evolution, 306, 8, doi:10.1007/s00606-020-01639-x, S2CID 211005874
13. Arias, S.; Terrazas, T.; Arreola-Nava, H.J.; Vazquez-Sanchez, M. & Cameron, K.M. (2005), "Phylogenetic relationships in Peniocereus (Cactaceae) inferred from plastid DNA sequence data", J. Plant Res., 118 (5): 317–328, doi:10.1007/s10265-005-0225-3, PMID 16143879, S2CID 5617876
14. Sánchez, D.; Arias, S. & Terrazas, T. (2014), "Phylogenetic relationships in Echinocereus (Cactaceae, Cactoideae)", Systematic Botany, 39 (4): 1183–1196, doi:10.1600/036364414X683831, S2CID 84505202
15. Gosline, Terrence M. (2009). "Galápagos Islands, Biology". In Gillespie, Rosemary G. & Clague, David A. (eds.). Encyclopedia of Islands. Berkeley: University of California Press. pp. 357–367. ISBN 978-0-520-25649-1.
16. Lendel, Anita (2013). South American Cacti in time and space: studies on the diversification of the tribe Cereeae, with particular focus on subtribe Trichocereinae (Cactaceae) (Dr. sc. nat. thesis). University of Zurich. doi:10.5167/UZH-93287. Retrieved 2023-10-10.
17. "Leptocereus", Taxonomy Browser, National Center for Biotechnology Information (NCBI), retrieved 2021-03-15
18. "Stenocereus griseus (Haw.) Buxb.", Plants of the World Online, Royal Botanic Gardens, Kew, retrieved 2021-03-16

• Echinocereeae
• Caryophyllales tribes