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Cactus
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===Classification=== {{main article|Taxonomy of the Cactaceae}} In 1984, it was decided that the Cactaceae Section of the [[International Organization for Succulent Plant Study]] should set up a working party, now called the International Cactaceae Systematics Group (ICSG), to produce consensus classifications down to the level of genera. Their system has been used as the basis of subsequent classifications. Detailed treatments published in the 21st century have divided the family into around 125β130 genera and 1,400β1,500 species, which are then arranged into a number of tribes and subfamilies.{{sfnp|ps=none|Anderson|2001|pp=99β103}}<ref name=Hunt06/><ref name=BarcYessHawk11/> The ICSG classification of the cactus family recognized four subfamilies, the largest of which was divided into nine tribes. The subfamilies were:{{sfnp|ps=none|Anderson|2001|pp=99β103}} {{multiple image |title=The four cactus subfamilies |total_width=320 | perrow = 2 |image1=Pereskia aculeata4 cropped.jpg |caption1=[[Pereskioideae]]: ''[[Pereskia aculeata]]'' |image2=Opuntia chlorotica 5 cropped.jpg |caption2=[[Opuntioideae]]: ''[[Opuntia chlorotica]]'' |image3=Maihuenia poeppigii 02 cropped.jpg |caption3=[[Maihuenioideae]]: ''[[Maihuenia poeppigii]]'' |image4=Warzenkaktus - Mammillaria elongata.JPG |caption4=Cactoideae: ''[[Mammillaria elongata]]'' }} * Subfamily Pereskioideae <small>K. Schumann</small> :The only genus in the ICSG classification was ''Pereskia''. It has features considered closest to the ancestors of the Cactaceae. Plants are trees or shrubs with leaves; their stems are smoothly round in cross section, rather than being ribbed or having tubercles.{{sfnp|ps=none|Anderson|2001|pp=99β103}} Two systems may be used in [[photosynthesis]], both the "normal" [[C3 carbon fixation|C<sub>3</sub> mechanism]] and [[crassulean acid metabolism]] (CAM)βan "advanced" feature of cacti and other [[Succulent plant|succulents]] that conserves water.<ref name=EdwaDono06/> :[[Molecular phylogenetics|Molecular phylogenetic]] studies showed that when broadly [[Circumscription (taxonomy)|circumscribed]], ''Pereskia'' was not [[monophyly|monophyletic]],<ref name=EdwaNyffDono05/><ref name=BarcYessHawk11/> and it has been split into three genera, ''[[Leuenbergeria]]'', ''[[Rhodocactus]]'' and a narrowly circumscribed ''[[Pereskia]]''.<ref name=Lode13/><ref name=AsaiMiya16/> ''Leuenbergeria'' is then placed on its own in a separate monogeneric subfamily, Leuenbergerioideae.<ref name=MaytMoli15/> * Subfamily [[Opuntioideae]] <small>K. Schumann</small> :Some 15 genera are included in this subfamily. They may have leaves when they are young, but these are lost later. Their stems are usually divided into distinct "joints" or "pads" ([[cladode]]s).{{sfnp|ps=none|Anderson|2001|pp=99β103}} Plants vary in size from the small cushions of ''[[Maihueniopsis]]''{{sfnp|ps=none|Anderson|2001|p=399}} to treelike species of ''Opuntia'', rising to {{convert|10|m|ft|abbr=on}} or more.{{sfnp|ps=none|Anderson|2001|p=485}} * Subfamily [[Maihuenioideae]] <small>P. Fearn</small> :The only genus is ''Maihuenia'', with two species, both of which form low-growing mats.{{sfnp|ps=none|Anderson|2001|p=398}} It has some features that are primitive within the cacti. Plants have leaves, and crassulean acid metabolism is wholly absent.{{sfnp|ps=none|Anderson|2001|pp=99β103}} * Subfamily [[Cactoideae]] :Divided into nine tribes, this is the largest subfamily, including all the "typical" cacti. Members are highly variable in habit, varying from tree-like to [[epiphyte|epiphytic]]. Leaves are normally absent, although sometimes very reduced leaves are produced by young plants. Stems are usually not divided into segments, and are ribbed or tuberculate. Two of the tribes, [[Hylocereeae]] and [[Rhipsalideae]], contain climbing or [[Epiphyte|epiphytic]] forms with a rather different appearance; their stems are flattened and may be divided into segments.{{sfnp|ps=none|Anderson|2001|pp=99β103}} [[Molecular phylogenetics|Molecular phylogenetic studies]] have supported the [[monophyly]] of three of these subfamilies (not Pereskioideae),<ref name=BarcYessHawk11/><ref name=EdwaNyffDono05/> but have not supported all of the tribes or even genera below this level; indeed, a 2011 study found only 39% of the genera in the subfamily Cactoideae sampled in the research were [[monophyletic]].<ref name=BarcYessHawk11/> Classification of the cacti currently remains uncertain and is likely to change.
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