Sempervivoideae

{{short description|Largest of 3 subfamilies in the flowering plant family Crassulaceae}} {{Automatic taxobox |image = Cambridge Botanical Gardens (4931401237).jpg |image_caption = ''Sempervivum tectorum'' |image_alt = Sempervivum tectorum, type species |taxon = Sempervivoideae |authority = Arn<ref>1832: Botany 112.</ref> |synonyms = * Sedeae * Sedoideae Berger |subdivision_ranks = Tribes |subdivision = * Telephieae 't Hart Ohba and Thieded ined. * Umbiliceae Meisn. * Semperviveae Dumort. * Aeonieae Thiede ined.{{efn|name=Reveal1|Reveal proposed the revised authority: Aeonieae Thiede ex Reveal{{sfn|Reveal|2012}}}} * Sedeae Fr. |type_genus = ''Sempervivum'' }}

'''Sempervivoideae''' is the largest of three subfamilies in the Saxifragales family Crassulaceae, with about 20–30 genera with succulent leaves. Unlike the two smaller subfamilies, it is distributed in temperate climates. The largest genus in this subfamily is ''Sedum'', with about 470 species. {{TOC limit|3}}

== Description ==

Succulent leaved plants. Unlike the other two smaller subfamilies, which are highly derived, Sempervivoideae retain the basic features of the family Crassulaceae.{{sfn|Thiede|Eggli|2007}} The Sempervivoideae contain many familiar horticultural plants, such as ''Sedum''.{{sfn|Mort et al|2010}}

== Taxonomy ==

Sempervivoideae has taxonomic priority over its synonym, Sedoideae,{{sfn|Gontcharova| Gontcharov|2007}} and is related to the other Crassulaceae subfamilies, as shown in this cladogram, although Messerschmid and colleagues (2020) state that these three subfamily clades are successive sisters, rather than Sempervivoideae being a direct sister only to Kalanchoideae.{{sfn|Messerschmid et al|2020}} {{cladogram | title=Cladogram I: Evolutionary lines in Crassulaceae{{sfn|Thiede|Eggli|2007}} | align=left | cladogram= {{clade |style="text-align:left; padding:2.5px; background:#eef"; width:500px; |label1= Crassulaceae |1={{clade |1=Crassuloideae |2={{clade |1=Kalanchoideae |2=Sempervivoideae }} }} }} }} {{Clear left}}

=== Subdivisions ===

Six clades within the Subfamily Sempervivoideae have been segregated into five tribes with about thirty genera.{{sfn|Thiede|Eggli|2007}} {| class="wikitable" border="1" |+ <big>Clades and tribes within Sempervivoideae</big> !Clade||Tribe |- | Acre||rowspan=2|Sedeae |- | Leucosedum |- | Aeonium||Aeonieae |- | Sempervivum||Semperviveae |- | Rhodiola||Umbiliceae |- |Hylotelephium||Telephieae |- |}

The composition of the five tribes are: (number of genera/species);{{sfn|Nikulin et al|2016}} * Telephieae (6/50) * Umbiliceae (4/100) * Semperviveae (2/75) * Aeonieae (3/70) * Sedeae (7/520)

and their relationship is shown in the cladogram: {{Cladogram|title= Cladogram II: Sempervivoideae tribes{{sfn|Thiede|Eggli|2007}} |align= left |cladogram={{clade| style=width:400px; |label1=Sempervivoideae |1={{clade | 1= Telephieae | 2= {{clade | 1= Umbilicieae | 2= {{clade | 1= Aeonieae | 2={{clade | 1= Semperviveae | 2= Sedeae }} }} }} }} }} }} {{Clear}} However, given the difficulty of establishing a stable classification, some authors prefer larger groupings, e.g.;{{sfn|Gontcharova |Gontcharov|2009 }} * Telephinae (Telephieae, Umbilicieae) * Sedinae (Semperviveae, Aeonieae, Sedeae)

Alternatively, Messerschmid and colleagues (2020), based on the largest analysis of subfamily taxa, propose the following clade structure (with tribes) and number of genera, species;{{sfn|Messerschmid et al|2020}} * Telephium (Telephieae, Umbiliceae) 9, 160 * Sempervivum (Semperviveae) 3, 60 * Aeonium (Aeonieae) 4, 67 (+ ''Sedum p.p.'' 8 sp.) * Leucosedum (Sedeae p.p.) 6,80 (+ ''Sedum p.p.'' 120 sp.) * Acre (Sedeae p.p.) 7, 205 (+ ''Sedum p.p.'' 345 sp.)

In this analysis, these clades and tribes were related as shown in this cladogram; {{Cladogram|title= {{vanchor|Cladogram III}}: Crassulaceae and Sempervivoideae clades and divisions{{sfn|Messerschmid et al|2020}} |align= left |caption=''Notes'': '''I''': Petrosedum clade is unplaced, due to discordance in analyses. In plastid phylogeny, it is one of the successive sister clades to the remainder of Sempervivoideae, without Telephium. But in Internal transcribed spacer (ITS) phylogeny Petrosedum it is sister to Aeonium, with Sempervivum/Jovibarba closest to Sedeae. Previously it had been placed within Sempervivum/Jovibarba. '''II''': Divergence times are in million years ago |cladogram={{clade| style=width:800px; |label1=Family Crassulaceae |sublabel1=100 |1={{clade |sublabel1=82 | 1= Subfamily Crassuloideae | 2= {{clade |sublabel1=71 |sublabel2=Subfamily Sempervivoideae | 1= Subfamily Kalanchoideae | 2= {{clade |sublabel1=66 |label1=Telephium clade

| 1= {{clade |sublabel1= |sublabel2=Tribe | 1=Tribe Telephiae |label2=Umbiliceae | 2={{clade |sublabel1=Clade |sublabel2=Clade | 1=Umbilicus | 2=Phedimus/Rhodiola }} }} | 2={{clade |sublabel1=Clade |sublabel2=Clade | 1= Sempervivum/Jovibarba (Tribe Semperviveae) | 2= Aeonium (Tribe Aeoniae) |label3=Sedeae |sublabel3=Tribe

| 3= {{clade |sublabel1=Clade |sublabel2=Clade | 1=Acre | 2=Leucosedum }} }} }} }} }} }} }} {{Clear}}

Semperviviae, Aeonieae and Sedeae are definable only by plesiomorphic features, with their genera all derived from within ''Sedum''. Segregates of ''Sedum'' occur in each of these, but lack sufficient features to allow them to be allocated to definitive genera.{{sfn|Thiede|Eggli|2007}}{{sfn|Mayuzumi|Ohba|2004}} {{Clear}}

{| |- |valign=top|[[File:Sedum spectabile - blossom top (aka).jpg|thumb|upright|left|''Hylotelephium spectabile''|alt=Hylotelephium spectabile, the showy stonecrop]] |valign=top|

=== Telephium clade ===

This monophyletic clade is sister to the remainder of the Sempervivoideae subfamily, and its three sublineages, Telephiae, Phedimus/Rhodiola and Umbilicus form a polytomy with the rest of the subfamily.{{sfn|Messerschmid et al|2020}}

==== Telephieae tribe ('t Hart) Ohba and Thiede ined. ====

The Telephieae{{efn|The name Telephieae may be illegitimate in this context, since it has been previously used as a synonym for Caryophyllaceae{{sfn|Reveal|2012}} }} genera consist of former infrageneric taxa of ''Sedum'', and are distributed primarily in East Asia, but with a few species found in Europe and N America (''Hylotelphium''). Defined by 5-merous flowers, free petals, flat, dentate leaves and tuberous roots or thickened rhizomes. Leaves usually in rosettes, except ''Hylotelephium'', petals often spotted, autumn flowering. The taxonomy remains unstable, with species ''Orostachys'' embedded within both ''Meterostachys'' and ''Hylotelephium''. Neither ''Hylotelphium'' nor ''Orostachys'' are considered monophyletic. But of the two sections of the genus, section ''Appendiculatae'' appears monophyletic and sister to ''Meterostachys''. ''Kungia'' is a segregate of a non-monophyletic ''Orostachys'' and appears to be a sister to ''Sinocrassula''.{{sfn|Messerschmid et al|2020}}

It is likely that diversification of the tribe occurred at the time of the formation of the Himalayas.{{sfn|Nikulin et al|2015}} Telephiae contains about 6 genera, with about 48–50 species, including:{{sfn|Gontcharova |Gontcharov|2007 }}{{sfn|Thiede|Eggli|2007}} {{div col|colwidth=24em}} * ''Hylotelephium'' H.Ohba c. 27–30 sp., N hemisphere * ''Kungia'' K.T.Fu 2 sp., SW China * ''Meterostachys'' Nakai c. 1 sp., E Asia * ''Orostachys'' Fisch. c. 5–15 sp, C-E Asia * ''Sinocrassula'' A.Berger c. 3–7 sp., Himalaya * ''Perrierosedum'' (A.Berger) H.Ohba ''incertae sedis'' c. 1 sp., Madagascar {{div col end}} |- |valign=top|[[File:Umbilicus intermedius 1.JPG|thumb|upright|left|''Umbilicus intermedius''|alt=Umbilicus intermedius, the common pennywort]] |valign=top|

==== Umbiliceae tribe Meisn. ====

The Umbeliceae occur mainly in temperate areas of Asia, flowering in spring to early summer, with about 4−5 genera and 100 species. The tribe consists of two subclades. The first, Phedimus/Rhodiola, includes ''Phedimus'', ''Rhodiola'' and ''Pseudosedum''. The latter has been included within ''Rhodiola'', or treated as sister to that genus. ''Aizopsis'' is included by some authors in ''Phedimus''. This subclade contains about 88 species found predominantly in Asia, but some species of ''Phedimus'' and ''Rhodiola'' in Europe and some ''Rhodiola'' in N America.{{sfn|Messerschmid et al|2020}}

The second is the genus ''Umbilicus'' with about 13 species, distributed in the Mediterranean, Macaronesia, SW Asia, Arabia and north to east Africa.{{sfn|Messerschmid et al|2020}}

These genera are among those that have been segregated from ''Sedum'', including:{{sfn|Thiede|Eggli|2007}} {{div col|colwidth=24em}} * ''Aizopsis'' Grulich * ''Phedimus'' Raf. c. 18–20 sp., Asia, Europe{{sfn|Ohba et al|2000}}{{sfn|Fu et al|2004}}{{sfn|ICN|2019}} * ''Pseudosedum'' A.Berger c. 10–12 sp., C Asia * ''Rhodiola'' L. c. 58–60 sp., Himalaya, Siberia, Europe, N America * ''Umbilicus'' DC. {{div col end}} |- |valign=top|[[File:Sedum rupestre 01 ies.jpg|thumb|upright|left|''Petrosedum rupestre''|alt=Petrosedum rupestre]] |valign=top|

=== Petrosedum clade ===

The phylogenetic relatpionships of the Petrosedum clade are not fully resolved, and hence is unplaced, due to discordance in molecular marker analyses. In plastid phylogeny, it is one of the successive sister clades to the remainder of Sempervivoideae, without Telephium. But in Internal transcribed spacer (ITS) phylogeny Petrosedum is sister to Aeonium, with Sempervivum/Jovibarba closest to Sedeae. Previously it had been placed within Sempervivum/Jovibarba. All species of ''Sedum'' series ''Rupestria'' A.Berger belong in this clade. This series has been elevated to genus rank as ''Petrosedum'', but ''Sedum'' series ''Nana'' also belongs to this segregated genus, or is at least sister to ''Petrosedum'' in this clade. ''Petrosedum'' as constituted from series ''Rupestria'' consists of 14 species from Europe, the Levant and N Africa. With the inclusion of series ''Nana'' from the Near East a further 9–12 species are included in the clade. As such the clade consists of a single genus;{{sfn|Messerschmid et al|2020}} * ''Petrosedum'' Grulich c. 23–26 sp. Euro-Mediterranean{{efn|''Petrosedum'' was classified as ''Sedum'' series ''Rupestria'' by some authors{{sfn|Nikulin et al|2016}}}}{{sfn|Gallo|2017}}

=== Sempervivum/Jovibarba clade (Semperviveae tribe Dumort.) ===

Semperviveae have acuminate leaves and polymerous flowers. Two genera, derived from within ''Sedum'', c. 275 species. Earlier treatments considered ''Jovibarba'' a section of ''Sempervivum''{{sfn|Thiede|Eggli|2007}} but has subsequently been demonstrated to be a separate genus.{{sfn|Mort et al|2010}} ''Petrosedum'' was originally also included in this clade. The two genera are oreophytes and occur predominantly in western Eurasian mountainous regions.{{sfn|Messerschmid et al|2020}} {{div col|colwidth=24em}} * ''Jovibarba'' (DC.) Opiz c. 2 sp. * ''Sempervivum'' L. c. 46 sp. {{div col end}} |- |valign=top|[[File:Aeonium aureum (≡ Greenovia. aurea) (Marianne Perdomo) 1.jpg|thumb|upright|left|''Aeonium aureum''|alt=Aeonium aureum]] |valign=top|

=== Aeonium clade (Aeonieae tribe) Thiede ined.{{efn|name=Reveal1}} ===

Aeonieae (also referred to as the GAMA clade, for ''Greenovia'', ''Aeonium'', ''Monanthes'' and ''Aichryson''),{{sfn|Soltis et al|2013}} whose distribution is primarily Macaronesia, have polymerous flowers, and include genera derived from within ''Sedum''. The clade includes four genera, including the monotypic ''Hypagophytum'' previously thought to be part of subfamily Crassuloideae, and about 64–70 species.{{sfn|Thiede|Eggli|2007}} It also includes about eight species of ''Sedum'' from three series (''Monanthoidea'', ''Caerulea'', ''Pubescens'') in northwest Africa and one species on Mediterranean islands. However ''Monanthes'' and ''Aeonium'' appear not to be monophyletic;{{sfn|Messerschmid et al|2020}} {{div col|colwidth=24em}} * ''Aeonium'' Webb & Berthel. (including ''Greenovia'' Webb) 39 sp. Macaronesia, Morocco, E Africa * ''Aichryson'' Webb & Berthel. 15 sp. Macaronesia * ''Hypagophytum'' A.Berger 1 sp. Ethiopia * ''Monanthes'' Haw. 10 sp. Macaronesia * ''Sedum'' L. 8 sp. NW Africa{{dubious|Taxonomic nonsense|reason=A genus cannot be in multiple tribes at once. If these are supposed to be various fragments of what used to be "Sedum", then referring to them collectively as "Sedum" is an error.|date=March 2026}} {{div col end}} |- |valign=top| |valign=top|

=== Sedeae Fr. ===

Sedeae is the largest tribe of these and has been considered as consisting of two sister clades, Leucosedum and Acre.{{sfn|Thiede|Eggli|2007}} It contains about 12 genera, mainly derived from within ''Sedum'', and 520 species. While the larger Acre clade is relatively robust, Leucosedum remains paraphyletic.{{sfn|Nikulin et al|2016}} In the largest study of this tribe, four polytomous clades are identified, of which the largest is Acre. Of the remaining three, ''Rosularia'', usually included in Leucosedum, is the smallest. In the remaining two clades, the species of ''Sedum'' belong to subgenus ''Gormania'', defined morphologically. Among the other genera, ''Dudleya'' and ''Sedella'' form a subclade.{{sfn|Thiede|Eggli|2007}}{{sfn|Messerschmid et al|2020}} |- |valign=top|[[File:2017-06-17-14.05.23 ZS PMax Dudleya caespitosa-1 - Flickr - John Rusk.jpg|thumb|upright|left|''Dudleya caespitosa''|alt=Dudleya caespitosa, the sea lettuce]] |valign=top|

==== Leucosedum clade ====

Genera derived from European and Mediterranean ''Sedum'' subg. ''Gormania'', with two western North American genera (''Dudleya'', ''Sedella'', which are sister genera). This clade has the largest topological discordance between the phylogenies defined by ITS and plastid markers. The genera included are:{{sfn|Nikulin et al|2016}}{{sfn|Messerschmid et al|2020}} {{colbegin|colwidth=24em}} * ''Rosularia'' (DC.) Stapf c. 20 sp., E Europe, Himalaya, Altai * ''Dudleya'' Britton & Rose N America * ''Sedella'' Britton & Rose N America * ''Afrovivella'' A.Berger * ''Pistorinia'' DC. Mediterranean * ''Prometheum'' (A.Berger) H.Ohba c. 8 sp., N Greece, SW Asia * (''Sedum'') {{colend}} |- |valign=top|[[File:Echeveria agavoides 01.jpg|thumb|upright|left|''Echeveria agavoides''|alt=Echeveria agavoides]] |valign=top|

==== Acre clade ====

The seven genera, including ''Sedum'' subgenus ''Sedum'', include 500 species. ''Villadia'', ''Echeveria'' and ''Graptopetalum'' are non-monophyletic:{{sfn|Messerschmid et al|2020}} {{colbegin|colwidth=24em}} * ''Echeveria'' DC. * ''Graptopetalum'' Rose * ''Lenophyllum'' Rose c. 7 sp. * ''Pachyphytum'' Link, Klotzsch & Otto * ''Thompsonella'' Britton & Rose * ''Villadia'' Rose * ''Sedum'' L. c. 470 sp. N hemisphere, S. America, N Africa{{dubious|Taxonomic nonsense|reason=A genus cannot be in multiple tribes at once. If these are supposed to be various fragments of what used to be Sedum, then referring to them collectively as "Sedum" is an error.|date=March 2026}} {{colend}} |} {{Clear}}

=== Genera === {{see also|List of Sempervivoideae genera}} Many of the genera in this subfamily have been considered non-monophyletic.{{sfn|Messerschmid et al|2020}} Other than the Sempervivum clade, ''Sedum'' has never formed a monophyletic group, but rather is scattered through the remaining clades, and thus is highly polyphyletic (or paraphyletic). This has been referred to as the "''Sedum'' problem".{{sfn|Messerschmid et al|2020}} Given the monophyly demonstrated for Aeonieae and Semperviveae (as quite distinct from Sedeae), it has been recommended that those species of ''Sedum'' originally found in those tribes, be removed from the genus and reassigned. This includes ''Sedum'' series ''Rupestria'' from Semperviveae, but collectively account for only a small fraction of the genus. While restricting Sedum to Sedeae simplifies the infrafamilial structure of the genus, its species remain distributed within both clades of this tribe.{{sfn|Nikulin et al|2016}}{{sfn|Gallo|2017}}{{sfn|Gallo|2017a}} ''Sedum'', with about 470 species, is by far the largest (and most problematic) genus within the subfamily, and the family Crassulaceae.{{sfn|Messerschmid et al|2020}}

=== Evolution and biogeography ===

There is no known fossil record of Crassulaceae. The Crassulaceae family evolved approximately 100 million years ago (mya) in southern Africa with the two most basal phylogenetic branches (Crassula, Kalanchoe) representing the predominantly southern African members.{{sfn|Hart|1997}}{{sfn|Thiede|Eggli|2007}} Divergence times are shown in Cladogram III. The family had a gradual evolution, with a basal split between Crassuloideae and the rest of the family (Kalanchoideae, Sempervivoideae) at 82 mya, and Sempervivoideae splitting from Kalanchoideae at 71 mya. The Sempervivoideae subsequently dispersed north to the Mediterranean region, and from there to Eastern Europe and Asia (Sempervivum and Leucosedum clades), with multiple groups spreading over the three continents of the Northern Hemisphere. The Telephium clade splitting from the rest of the subfamily at 66 mya. This was followed by the Petrosedum and Aeonium clades at 56 mya and Sempervivum/Jovibarda at 52 mya. The remaining two clades, constituting Sedeae (Leucosedum and Acre) separating from each other at 48 mya. Two lineages from the European Crassulaceae eventually dispersed to North America and underwent subsequent diversification. The Aeonium clade dispersed from northern Africa to adjacent Macaronesia.{{sfn|Mort et al|2001}}{{sfn|Messerschmid et al|2020}}

== Notes == {{notelist}}

== References == {{Reflist|20em}}

== Bibliography == {{refbegin|30em}} === Books === * {{cite book|editor-last=Eggli|editor-first=Urs|title=Illustrated Handbook of Succulent Plants: Crassulaceae|url=https://books.google.com/books?id=nU7mCAAAQBAJ|date=2003|publisher=Springer Science & Business Media|doi=10.1007/978-3-642-55874-0|isbn=978-3-642-55874-0}} * {{cite book |last1=Thiede |first1=J |last2=Eggli |first2=U |editor-last=Kubitzki |editor-first=Klaus |editor-link=Klaus Kubitzki |title=Berberidopsidales, Buxales, Crossosomatales, Fabales p.p., Geraniales, Gunnerales, Myrtales p.p., Proteales, Saxifragales, Vitales, Zygophyllales, Clusiaceae Alliance, Passifloraceae Alliance, Dilleniaceae, Huaceae, Picramniaceae, Sabiaceae |year=2007 |chapter=Crassulaceae |pages=83–119 |publisher=Springer |url=https://books.google.com/books?id=PdSL7jBNX9EC |isbn=978-3-540-32214-6 }} ''([https://www.researchgate.net/publication/227205999_Crassulaceae full text at]'' ResearchGate)

=== Articles === * {{cite journal |last1=Carrillo-Reyes |first1=Pablo |last2=Sosa |first2=Victoria |last3=Mort |first3=Mark E. |title=Molecular phylogeny of the Acre clade (Crassulaceae): Dealing with the lack of definitions for Echeveria and Sedum |journal=Molecular Phylogenetics and Evolution |date=October 2009 |volume=53 |issue=1 |pages=267–276 |doi=10.1016/j.ympev.2009.05.022 |pmid=19482091 |bibcode=2009MolPE..53..267C |url=http://www.crassulaceae.com/crassulaceae.com/botanik/pflanzen/scans/gnr1630/1340-5.pdf |ref={{harvid|Carillo-Reyes et al|2009}} }} * {{cite journal |last1=Gallo |first1=Lorenzo |title=Towards a review of the genus ''Petrosedum'' (Crassulaceae): Taxonomic and nomenclatural notes on Iberian taxa |journal=Webbia |date=24 August 2017a |volume=72 |issue=2 |pages=207–216 |doi=10.1080/00837792.2017.1363978|bibcode=2017Webbi..72..207G |doi-access=free }} * {{cite journal |last1=Gallo |first1=Lorenzo |title=Nomenclatural novelties in ''Petrosedum'' (Crassulaceae) |journal=Phytotaxa |date=9 May 2017 |volume=306 |issue=2 |page=169 |doi=10.11646/phytotaxa.306.2.8|bibcode=2017Phytx.306..169G }} * {{cite journal |last1=Gontcharova |first1=S. B. |last2=Artyukova |first2=E. V. |last3=Gontcharov |first3=A. A. |title=Phylogenetic relationships among members of the subfamily Sedoideae (Crassulaceae) inferred from the ITS region sequences of nuclear rDNA |journal=Russian Journal of Genetics |date=June 2006 |volume=42 |issue=6 |pages=654–661 |doi=10.1134/S102279540606010X |url=http://biosoil.ru/files/00002900.pdf |ref={{harvid|Gontcharova et al|2006}} |archive-date=2017-08-09 |access-date=2019-09-27 |archive-url=https://web.archive.org/web/20170809055756/http://www.biosoil.ru/files/00002900.pdf |url-status=dead }} * {{cite journal |last1=Gontcharova |first1=Svetlana B. |last2=Gontcharov |first2=Andrey A. |title=Molecular Phylogenetics of Crassulaceae |journal=Genes, Genomes and Genomics |date=2007 |volume=1 |issue=1 |pages=40–46 |url=http://www.globalsciencebooks.info/Online/GSBOnline/images/0706/GGG_1(1)/GGG_1(1)40-46o.pdf }} * {{cite journal |last1=Gontcharova |first1=S. B. |last2=Gontcharov |first2=A. A. |title=Molecular phylogeny and systematics of flowering plants of the family Crassulaceae DC |journal=Molecular Biology |date=11 October 2009 |volume=43 |issue=5 |pages=794–803 |doi=10.1134/S0026893309050112}} * {{cite journal |last1=Grulich |first1=Vit |title=Generic division of Sedoideae in Europe and the adjacent regions |journal=Preslia |date=1984 |volume=56 |pages=29–45 |url=http://www.preslia.cz/archive/Preslia_56_1984_29-45.pdf }} * {{cite journal |last1=Hart |first1=H. 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=== Websites === * {{cite web |title=International Crassulaceae Network |url=http://www.crassulaceae.ch/de/home |access-date=5 September 2019 |ref={{harvid|ICN|2019}} }} * {{cite web |last1=Fu |first1=Kunjun |last2=Ohba |first2=Hideaki |last3=Gilbert |first3=Michael G. |title=Crassulaceae Candolle |url=http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=10225 |page=202 |access-date=24 August 2019 |year=2004 |ref={{harvid|Fu et al|2004}} }}, in Flora of China online vol. 8 {{refend}}

== External links == {{Commons}}{{Wikispecies}}

Category:Crassulaceae Category:Eudicot subfamilies