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{{Automatic taxobox
|image = Cambridge Botanical Gardens (4931401237).jpg
|image_caption = ''[[Sempervivum tectorum]]''
|image_alt = Sempervivum tectorum, type species
|taxon = Sempervivoideae
|authority = [[George Arnott Walker-Arnott|Arn]]<ref>1832: Botany 112.</ref>
|synonyms =
* Sedeae
* Sedoideae <small>[[A.Berger|Berger]]</small>
|subdivision_ranks = Tribes
|subdivision =
* Telephieae <small>'t Hart Ohba and Thieded [[ined.]]</small>
* Umbiliceae <small>[[Meisn.]]</small>
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'''Sempervivoideae''' is the largest of three [[subfamilies]] in the [[Saxifragales]] [[family (biology)|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 ==
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== Taxonomy ==
Sempervivoideae has taxonomic priority over its [[synonym (taxonomy)|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
{{cladogram
| title=Cladogram
| align=left
| cladogram=
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|1=Crassuloideae
|2={{clade
|1=
|2=Sempervivoideae
}}
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{{Clear left}}
=== Subdivisions
Six [[clades]] within
{| class="wikitable" border="1"
|+ <big>Clades and tribes within Sempervivoideae</big>
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and their relationship is shown in the cladogram:
{{Cladogram|title= Cladogram
|align= left
|cladogram={{clade| style=width:{{{width|400}}}px;
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| 2={{clade
| 1= Semperviveae
| 2= Sedeae
}}
}}
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* 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
|align= left
|caption=
|cladogram={{clade| style=width:{{{width|800}}}px;
|label1=Family Crassulaceae
|sublabel1=
|1={{clade
|sublabel1=
| 1= Subfamily Crassuloideae
| 2= {{clade
|sublabel1=
|sublabel2=Subfamily Sempervivoideae
| 1= Subfamily Kalanchoideae
| 2= {{clade
|sublabel1=
|label1=Telephium clade
| 1= {{clade
|sublabel1=
|sublabel2=Tribe
| 1=Tribe Telephiae
|label2=Umbiliceae
| 2={{clade
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}}
{{Clear}}
Semperviviae, Aeonieae and Sedeae are definable only by [[plesiomorphic]] features, with their genera all derived from within ''Sedum''. [[Segregate (taxonomy)|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}}
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The Telephieae{{efn|The name Telephieae may be [[Nomen illegitimum|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 leaf|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 [[section (biology)|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]]'' <small>H.Ohba</small> c. 27–30 sp., N hemisphere
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==== Umbiliceae tribe <small>[[Meisn.]]</small> ====
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
These genera are among those that have been [[segregate (taxonomy)|segregated]] from ''Sedum'', including:{{sfn|Thiede|Eggli|2007}}
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* ''[[Petrosedum]]'' <small>Grulich</small> 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}}
|-
|valign=top|[[File:Sempervivum ciliosum 01.jpg|thumb|upright|left|''[[Sempervivum tectorum]]''|alt=Sempervivum tectorum, the common houseleek]]
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=== Sempervivum/Jovibarba clade (Semperviveae tribe <small>[[Dumort.]]</small>) ===
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]]'' <small>(DC.) Opiz</small> c. 2 sp.
* ''[[Sempervivum]]'' <small>L.</small> c. 46 sp.
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* ''[[Hypagophytum]]'' <small>A.Berger</small> 1 sp. Ethiopia
* ''[[Monanthes]]'' <small>Haw.</small> 10 sp. Macaronesia
* ''[[Sedum]]'' <small>L.</small> 8 sp. NW Africa
{{div col end}}
|-
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==== 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]]'' <small>(DC.) Stapf</small> c. 20 sp., E Europe, Himalaya, Altai
* ''[[Dudleya]]'' <small>Britton & Rose</small> N America
* ''[[Sedella (plant)|Sedella]]'' <small>Britton & Rose</small> N America
▲* ''[[Afrovivella]]'' <small>A.Berger</small>
* ''[[Pistorinia]]'' <small>DC.</small> Mediterranean
* ''[[Prometheum]]'' <small>(A.Berger) H.Ohba</small> c. 8 sp., N Greece, SW Asia
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==== Acre clade ====
The seven genera,
{{colbegin|colwidth=24em}}
* ''[[Echeveria]]'' <small>DC.</small>
* ''[[Graptopetalum]]'' <small>Rose</small>
* ''[[Lenophyllum]]'' <small>Rose</small> c. 7 sp.
* ''[[Pachyphytum]]'' <small>Link, Klotzsch & Otto</small>
* ''[[Thompsonella]]'' <small>Britton & Rose</small>
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{{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 ([[myr|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|Cladogram III]]. The family had a gradual evolution, with a [[Basal (phylogenetics)|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 ==
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=== 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|url=https://books.google.com/books?id=PdSL7jBNX9EC |isbn=978-3540322146}} ''([https://www.researchgate.net/publication/227205999_Crassulaceae full text at]'' [[
=== 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 |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}}
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* {{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. 't |title=The evolution of the Sedum acre group (Crassulaceae) |journal=Bocconea |date=1995 |volume=5 |pages=119–128 |url=http://www.herbmedit.org/bocconea/5-119.pdf}}
* {{cite journal |last= Hart |first=H.'t |year=1997 |title=Diversity within Mediterranean Crassulaceae |url=https://idus.us.es/xmlui/bitstream/handle/11441/63144/Diversity%20Hart.pdf?sequence=1|journal=Lagascalia |volume=1 |issue=2 |pages=93–100}}
* {{cite journal |last1=Lim |first1=Mi Sang |last2=Choi |first2=Sun Hee |title=Estimation of Phylogeny of Nineteen Sedoideae Species Cultivated in Korea Inferred from Chloroplast DNA Analysis |journal=The Horticulture Journal |date=2018 |volume=87 |issue=1 |pages=132–139 |doi=10.2503/hortj.OKD-087|doi-access=free }}
* {{cite journal |last1=Mayuzumi |first1=Shinzo |last2=Ohba |first2=Hideaki |title=The Phylogenetic Position of Eastern Asian Sedoideae (Crassulaceae) Inferred from Chloroplast and Nuclear DNA Sequences |journal=Systematic Botany |date=2004 |volume=29 |issue=3 |pages=587–598 |issn=0363-6445|jstor=25063994 |doi=10.1600/0363644041744329 }}
* {{cite journal |last1=Messerschmid |first1=Thibaud F.E. |last2=Klein |first2=Johannes T. |last3=Kadereit |first3=Gudrun |last4=Kadereit |first4=Joachim W. |title=Linnaeus's folly – phylogeny, evolution and classification of Sedum (Crassulaceae) and Crassness subfamily Sempervivoideae |journal=[[Taxon (journal)|Taxon]] |date=4 September 2020 |pages=tax.12316 |doi=10.1002/tax.12316|ref={{harvid|Messerschmid et al|2020}}|doi-access=free }}
* {{cite journal |last1=Mort |first1=Mark E. |last2=Soltis |first2=Douglas E. |last3=Soltis |first3=Pamela S. |last4=Francisco-Ortega |first4=Javier |last5=Santos-Guerra |first5=Arnoldo |authorlink2=Douglas E. Soltis|authorlink3=Pamela Soltis| title=Phylogenetic relationships and evolution of Crassulaceae inferred from matK sequence data |journal=[[American Journal of Botany]] |date=January 2001 |volume=88 |issue=1 |pages=76–91 |doi=10.2307/2657129|jstor=2657129|pmid=11159129 |ref={{harvid|Mort et al|2001}} |doi-access=free }}
* {{cite journal |last1=Mort |first1=Mark E |last2=O'Leary |first2=T. Ryan |last3=Carrillo-Reyes |first3=Pablo |display-authors=etal|title=Phylogeny and evolution of Crassulaceae: Past, present, and future |journal=Biodiversity & Ecology |date=December 2010 |volume=3 |pages=69–86 |url=https://www.researchgate.net/publication/248701315|ref={{harvid|Mort et al|2010}}}}
* {{cite journal |last1=Nikulin |first1=Arthur |last2=Vyacheslav Nikulin |first2=Vyacheslav |last3=Gontcharov |first3=Andrey |title=To the question of phylogenetic structure of the tribe Telephieae (Sempervivoideae, Crassulaceae) based on ITS rDNA sequence comparisons |journal=Botanicheskii Zhurnal |date=October 2015 |volume=100 |issue=10 |pages=1030–1040 |url=https://www.researchgate.net/publication/311213406 |language=ru|ref={{harvid|Nikulin et al|2015}}}}
* {{cite journal |last1=Nikulin |first1=Vyacheslav Yu. |last2=Gontcharova |first2=Svetlana B. |last3=Stephenson |first3=Ray |last4=Gontcharov |first4=Andrey A. |title=Phylogenetic relationships between Sedum L. and related genera (Crassulaceae) based on ITS rDNA sequence comparisons |journal=Flora |date=September 2016 |volume=224 |pages=218–229 |doi=10.1016/j.flora.2016.08.003|ref={{harvid|Nikulin et al|2016}}}}
* {{cite journal |last1=Ohba |first1=Hideaki |last2=Bartholomew |first2=Bruce M |last3=Turland |first3=Nicholas J |last4=Kunjun |first4=Fu |title=New Combinations in Phedimus (Crassulaceae) |journal=Novon |date=2000 |volume=10 |issue=4 |pages=400–402 |url=
* {{cite journal |last1=Ohba |first1=H |title=Generic and infrageneric classification of the old world sedoideae crassulaceae |journal=Journal of the Faculty of Science University of Tokyo Section III Botany
* {{cite journal |last1=Reveal |first1=James L |authorlink=James L. Reveal|title=An outline of a classification scheme for extant flowering plants |journal=Phytoneuron |date=2012 |volume=37 |pages=1–221 |url=https://www.phytoneuron.net/PhytoN-Magnoliidae.pdf}}
* {{cite journal |last1=Soltis |first1=D. E. |authorlink1=Douglas E. Soltis |last2=Mort |first2=M. E. |last3=Latvis |first3=M. |last4=Mavrodiev |first4=E. V. |last5=O'Meara |first5=B. C. |last6=Soltis |first6=P. S. |authorlink6 = Pamela S. Soltis|last7=Burleigh |first7=J. G. |last8=Rubio de Casas |first8=R. |title=Phylogenetic relationships and character evolution analysis of Saxifragales using a supermatrix approach |journal=[[American Journal of Botany]] |date=29 April 2013 |volume=100 |issue=5 |pages=916–929 |doi=10.3732/ajb.1300044|pmid=23629845 |ref={{harvid|Soltis et al|2013}}}}
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<!--- Categories --->
[[Category:Crassulaceae]]
[[Category:
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