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Opuntioideae

Opuntioideae is a subfamily of the cactus family, Cactaceae. It contains 15 genera divided into five tribes. The subfamily encompasses roughly 220–250 species, and is geographically distributed throughout the New World from Canada, to Argentina.[2] Members of this subfamily have diverse habits, including small geophytes, hemispherical cushions, shrubs, trees, and columnar cacti consisting of indeterminate branches or determinate terete or spherical segments.[3]

Description edit

Synapomorphies of Opuntioideae include small deciduous, barbed spines called glochids born on areoles and a bony aril surrounding a campylotropous ovule (inverted and curved, such that the micropyle almost meets the funiculus).[4] Other prominent morphological characters for this subfamily are presence of cylindrical, caducous leaves that tend to be shed by maturity and the sectioning of the stem into joints or pads known as cladodes.

Opuntioideae are unique among cacti for lacking in the stem a thick cortex, an extensive system of cortical bundles, collapsible cortical cells, and medullary bundles. Typically, the epidermis consists of a single layer of irregularly shaped cells, a cuticle at least 1-2 microns thick, and long, uniseriate trichomes in the areoles. Opuntioideae have a hypodermis of at least one layer, very thick walls, and druses (aggregations of calcium oxalate crystals), and their cortical cells have enlarged nuclei; the reason for this is unknown. They also possess mucilage cells.

Notably, their lack of collapsible cortical cells, ribs, and tubercles mean that they cannot absorb water or transfer it intercellularly as easily as the other cacti, so this may place evolutionary constraints on the aridity of habitats and maximum adult size. One adaptation around this problem is the evolution of flattened cladodes that allow opuntioids to swell up with water, increasing in volume without an increase in surface area risking water loss. Opuntioids also lack fiber caps to their phloem bundles, which in other cacti protect against sucking insects and stiffen developing internodes.[5]

Tribes and genera edit

Tribes and genera of the subfamily Opuntioideae include:[6]

 
Austrocylindropuntia subulata

Cylindropuntieae – round stems

 
Cylindropuntia bigelovii

Opuntieae – flattened stems


Tephrocacteae – round stems

Intergeneric Hybrids edit

  • × Opulea M.H.J.van der Meer (Consolea × Opuntia)

Systematics edit

Current research shows conflicting arguments over the phylogenetic accuracy of the tribes and genera listed above. A major challenge in Opuntioideae classification is that the subfamily is known to hybridize (particularly the Opuntieae tribe), which further complicates how to define at the species level.[7] In addition, the genera within each tribe exhibits variation in morphology, which makes using genetic analysis more important in determining relationships, since defining physical characteristics may be unseen in some genera.[8]

In 2009, a study by Griffith and Porter, based on ribosomal intergenic spacer analysis defined four tribes of Opuntioideae:[4]

  1. Core Maihueniopsis - which was shown as monophyletic through genetic analysis
  2. Pterocactus - defined by a winged seed
  3. Terete-stemmed - defined by cylinder-shaped stems
  4. Flat-stemmed - defined by flat stems

However, this classification has been questioned by additional research that did not yield the same systematic results. Specifically, there was differences in yielded relationships based on what genetic sequences and analysis was being utilized.[3][9]

In 2010, it was proposed by Nyffeler and Eggli, as part of a larger overhaul of Cactaceae systematics, to only recognize Cylindropuntieae and Opuntieae as true tribes of Opuntioideae, since they are the tribes generally agreed upon to be monophyletic. It was proposed to include all other genera into one polyphyletic basal group until further research concludes more definitive answers to their phylogeny.[10]

See also edit

References edit

  1. ^ . Germplasm Resources Information Network. United States Department of Agriculture. 2004-02-13. Archived from the original on 2012-08-25. Retrieved 2011-12-05.
  2. ^ Anderson, Edward F. (2001). The Cactus Family. Portland, OR: Timber Press. ISBN 0881924989.
  3. ^ a b Ritz, C.M.; Reiker, J.; Charles, G.; Hoxey, P.; Hunt, D.; Lowry, M.; Stuppy, W.; Taylor, N. (2012). "Molecular phylogeny and character evolution in terete-stemmed Andean opuntias (Cactaceae−Opuntioideae)". Molecular Phylogenetics and Evolution. 65 (2): 668–681. doi:10.1016/j.ympev.2012.07.027. PMID 22877645.
  4. ^ a b Griffith, M. Patrick; Porter, J. Mark (2009-01-01). "Phylogeny of Opuntioideae (Cactaceae)". International Journal of Plant Sciences. 170 (1): 107–116. doi:10.1086/593048. ISSN 1058-5893.
  5. ^ Mauseth, James D. (2005-12-01). "Anatomical features, other than wood, in subfamily opuntioideae (cactaceae)". Haseltonia. 11: 113–125. doi:10.2985/1070-0048(2005)11[113:afotwi]2.0.co;2. ISSN 1070-0048.
  6. ^ . Germplasm Resources Information Network. United States Department of Agriculture. Archived from the original on 2015-09-24. Retrieved 2011-12-05.
  7. ^ Majure, Lucas; Judd, Walter; Soltis, Pam; Soltis, Doug (2012-02-14). "Cytogeography of the Humifusa clade of Opuntia s.s. Mill. 1754 (Cactaceae, Opuntioideae, Opuntieae): correlations with pleistocene refugia and morphological traits in a polyploid complex". Comparative Cytogenetics. 6 (1): 53–77. doi:10.3897/compcytogen.v6i1.2523. ISSN 1993-078X. PMC 3833768. PMID 24260652.
  8. ^ Bárcenas, Rolando T. (August 2016). "A molecular phylogenetic approach to the systematics of Cylindropuntieae (Opuntioideae, Cactaceae)". Cladistics. 32 (4): 351–359. doi:10.1111/cla.12135.
  9. ^ Hernandez-Hernandez, T.; Hernandez, H. M.; De-Nova, J. A.; Puente, R.; Eguiarte, L. E.; Magallon, S. (2010-12-23). "Phylogenetic relationships and evolution of growth form in Cactaceae (Caryophyllales, Eudicotyledoneae)". American Journal of Botany. 98 (1): 44–61. doi:10.3732/ajb.1000129. ISSN 0002-9122. PMID 21613084.
  10. ^ Nyffeler, Retro, Eggli, Urs (2010). "A Farewell to Dated Ideas and Concepts: Molecular Phylogenetics and a Revised Suprageneric Classification of the family Cactaceae" (PDF). Schumannia. 6: 109–149 – via uzh.ch.{{cite journal}}: CS1 maint: multiple names: authors list (link)

opuntioideae, subfamily, cactus, family, cactaceae, contains, genera, divided, into, five, tribes, subfamily, encompasses, roughly, species, geographically, distributed, throughout, world, from, canada, argentina, members, this, subfamily, have, diverse, habit. Opuntioideae is a subfamily of the cactus family Cactaceae It contains 15 genera divided into five tribes The subfamily encompasses roughly 220 250 species and is geographically distributed throughout the New World from Canada to Argentina 2 Members of this subfamily have diverse habits including small geophytes hemispherical cushions shrubs trees and columnar cacti consisting of indeterminate branches or determinate terete or spherical segments 3 OpuntioideaeCylindropuntia ramosissimaScientific classificationKingdom PlantaeClade TracheophytesClade AngiospermsClade EudicotsOrder CaryophyllalesFamily CactaceaeSubfamily OpuntioideaeBurnettTribesCylindropuntieaeOpuntieaePterocacteaeTephrocacteae 1 Contents 1 Description 2 Tribes and genera 3 Intergeneric Hybrids 4 Systematics 5 See also 6 ReferencesDescription editSynapomorphies of Opuntioideae include small deciduous barbed spines called glochids born on areoles and a bony aril surrounding a campylotropous ovule inverted and curved such that the micropyle almost meets the funiculus 4 Other prominent morphological characters for this subfamily are presence of cylindrical caducous leaves that tend to be shed by maturity and the sectioning of the stem into joints or pads known as cladodes Opuntioideae are unique among cacti for lacking in the stem a thick cortex an extensive system of cortical bundles collapsible cortical cells and medullary bundles Typically the epidermis consists of a single layer of irregularly shaped cells a cuticle at least 1 2 microns thick and long uniseriate trichomes in the areoles Opuntioideae have a hypodermis of at least one layer very thick walls and druses aggregations of calcium oxalate crystals and their cortical cells have enlarged nuclei the reason for this is unknown They also possess mucilage cells Notably their lack of collapsible cortical cells ribs and tubercles mean that they cannot absorb water or transfer it intercellularly as easily as the other cacti so this may place evolutionary constraints on the aridity of habitats and maximum adult size One adaptation around this problem is the evolution of flattened cladodes that allow opuntioids to swell up with water increasing in volume without an increase in surface area risking water loss Opuntioids also lack fiber caps to their phloem bundles which in other cacti protect against sucking insects and stiffen developing internodes 5 Tribes and genera editTribes and genera of the subfamily Opuntioideae include 6 nbsp Austrocylindropuntia subulataCylindropuntieae round stems nbsp Cylindropuntia bigeloviiCylindropuntia Engelm F M Knuth Grusonia Rchb f ex Britton amp Rose Pereskiopsis Britton amp Rose Quiabentia Britton amp RoseOpuntieae flattened stems Brasiliopuntia K Schum A Berger Consolea Lem Miqueliopuntia Fric ex F Ritter Opuntia Mill Salmonopuntia P V Heath Tacinga Britton amp Rose Airampoa FricTephrocacteae round stems Austrocylindropuntia Backeb Cumulopuntia F Ritter Maihueniopsis Speg Tephrocactus Lem Pterocactus K Schum Punotia D R HuntIntergeneric Hybrids edit Opulea M H J van der Meer Consolea Opuntia Systematics editSee also Taxonomy of the Cactaceae Current research shows conflicting arguments over the phylogenetic accuracy of the tribes and genera listed above A major challenge in Opuntioideae classification is that the subfamily is known to hybridize particularly the Opuntieae tribe which further complicates how to define at the species level 7 In addition the genera within each tribe exhibits variation in morphology which makes using genetic analysis more important in determining relationships since defining physical characteristics may be unseen in some genera 8 In 2009 a study by Griffith and Porter based on ribosomal intergenic spacer analysis defined four tribes of Opuntioideae 4 Core Maihueniopsis which was shown as monophyletic through genetic analysis Pterocactus defined by a winged seed Terete stemmed defined by cylinder shaped stems Flat stemmed defined by flat stemsHowever this classification has been questioned by additional research that did not yield the same systematic results Specifically there was differences in yielded relationships based on what genetic sequences and analysis was being utilized 3 9 In 2010 it was proposed by Nyffeler and Eggli as part of a larger overhaul of Cactaceae systematics to only recognize Cylindropuntieae and Opuntieae as true tribes of Opuntioideae since they are the tribes generally agreed upon to be monophyletic It was proposed to include all other genera into one polyphyletic basal group until further research concludes more definitive answers to their phylogeny 10 See also editTaxonomy of the CactaceaeReferences edit 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 2011 12 05 Anderson Edward F 2001 The Cactus Family Portland OR Timber Press ISBN 0881924989 a b Ritz C M Reiker J Charles G Hoxey P Hunt D Lowry M Stuppy W Taylor N 2012 Molecular phylogeny and character evolution in terete stemmed Andean opuntias Cactaceae Opuntioideae Molecular Phylogenetics and Evolution 65 2 668 681 doi 10 1016 j ympev 2012 07 027 PMID 22877645 a b Griffith M Patrick Porter J Mark 2009 01 01 Phylogeny of Opuntioideae Cactaceae International Journal of Plant Sciences 170 1 107 116 doi 10 1086 593048 ISSN 1058 5893 Mauseth James D 2005 12 01 Anatomical features other than wood in subfamily opuntioideae cactaceae Haseltonia 11 113 125 doi 10 2985 1070 0048 2005 11 113 afotwi 2 0 co 2 ISSN 1070 0048 GRIN Genera of Cactaceae subfam Opuntioideae Germplasm Resources Information Network United States Department of Agriculture Archived from the original on 2015 09 24 Retrieved 2011 12 05 Majure Lucas Judd Walter Soltis Pam Soltis Doug 2012 02 14 Cytogeography of the Humifusa clade of Opuntia s s Mill 1754 Cactaceae Opuntioideae Opuntieae correlations with pleistocene refugia and morphological traits in a polyploid complex Comparative Cytogenetics 6 1 53 77 doi 10 3897 compcytogen v6i1 2523 ISSN 1993 078X PMC 3833768 PMID 24260652 Barcenas Rolando T August 2016 A molecular phylogenetic approach to the systematics of Cylindropuntieae Opuntioideae Cactaceae Cladistics 32 4 351 359 doi 10 1111 cla 12135 Hernandez Hernandez T Hernandez H M De Nova J A Puente R Eguiarte L E Magallon S 2010 12 23 Phylogenetic relationships and evolution of growth form in Cactaceae Caryophyllales Eudicotyledoneae American Journal of Botany 98 1 44 61 doi 10 3732 ajb 1000129 ISSN 0002 9122 PMID 21613084 Nyffeler Retro Eggli Urs 2010 A Farewell to Dated Ideas and Concepts Molecular Phylogenetics and a Revised Suprageneric Classification of the family Cactaceae PDF Schumannia 6 109 149 via uzh ch a href Template Cite journal html title Template Cite journal cite journal a CS1 maint multiple names authors list link Retrieved from https en wikipedia org w index php title Opuntioideae amp oldid 1188197369, wikipedia, wiki, book, books, library,

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