Crassulaceae
Crassulaceae | |
---|---|
Crassula perfoliata Type species | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Eudicots |
Order: | Saxifragales |
Family: | Crassulaceae J.St.-Hil.[1] |
Type genus | |
Crassula | |
Subfamilies | |
| |
Synonyms | |
Sempervivae [Sempervivaceae] Jussieu[2] |
The Crassulaceae (
Crassulaceae are mainly perennial and have huge economic importance, internationally, as collectible, indoor and
Description
General: Crassulaceae is a family of
Vegetative:
Reproductive: The
The flowers are often
However, a number of genera (e.g. Sempervivum, Aeonium) are polymerous (3-32), have basally fused or partially fused corolla segments, where the petals may form a corolla tube of varying length (e.g. Kalanchoe, Cotyledon), or have only a single whorl of 5 stamens (e.g. Crassula, Tillaea),[11] while Sedum includes much of the morphological diversity within the family as a whole.[9] Although the typical number of floral plants is four or five, a number of genera, such as Sempervivum and Jovibarba, demonstrate polymery (at least ten or greater parts).[9]
Genome
Phytochemistry
Crassulacean acid metabolism (CAM photosynthesis) is named after the family, because the pathway was first discovered in crassulacean plants. It is one of the few families that still has CAM as an active, photosynthetic pathway, and is unique in which all its members are known to possess CAM.[3]
Taxonomy
Originally described by
One of the most complete treatments was
Molecular phylogenetics has shown that morphological characters and chromosome numbers are so labile in the family, with rampant polyploidy and aneuploidy, that they cannot be used reliably to infer evolution, even at low taxonomic levels, with few exceptions. For instance Prometheum and Rosularia have been segregated from Sedum by their basic chromosome numbers.[4][5][18]
Crassulaceae is a medium size
Phylogeny
Crassulaceae has been considered a part of the order
Cladogram of Saxifragales showing the evolutionary affinities of Crassulaceae | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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This Monogeneric families are represented by genus names, with family in parentheses. Unless otherwise stated, support for most nodes is maximal and equal to 100%.
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Biogeography and evolution
Crassulaceae evolved approximately 100–60 million years ago in southern Africa with the two most basal phylogenetic branches (Crassula, Kalanchoe) representing the predominantly southern African members.
Distinct centers of speciation developed in Macaronesia (Aeonium clade), Mexico (Sedum and Echeverioideae in clade 7), and southeastern Asia (Sedum sarmentosum, and S. morissonensis in Acre clade). On arrival in the Northern hemisphere the Sempervivoideae reached its greatest diversity. Conversely, few representatives of the Crassulaceae occur in South America and Australia.[5][9] Sedum species are found in most of these regions, generally grouped with genera endemic to that region. For instance the North African S. jaccardianum and S. modestum (Aeonium) are a sister group to the endemic Macaronesian species in that clade.[9] The Macaronesian archipelago appears to have been reached by Crassulaceae at least three times. Once by the ancestor of Aeonium and Monanthes, most likely from the Western Mediterranean region, with the closest extant relatives of these two genera (Sedum caeruleum, S. pubescens), coming from this region (Aeonium clade). The second migration was by an ancestor of a clade of three Sedum species (S. nudum, S. lancerotense and S. fusiforme (Acre clade)), which appear to have originated in Mexico. The third occurrence likely involved the ancestor of a lineage within the genus Umbilicus (Rhodiola clade). The Macronesian flora include three genera from the Sempervivoideae, Aeonium, Aichryson and Monanthes (Aeonium clade), together with several Sedum spp. and one species of Umbilicus (Rhodiola). North America was reached at least twice, once by an ancestor of Parvisedum and Dudleya, and once by a subclade of Acre. For a mapping of morphological features and biogeography on the phylogenetic tree, see Mort et al 2001 Fig. 3.[5][9]
Chromosome numbers have played a limited role in elucidating evolution, but suggest a core of x=8, with subsequent polyploidy. For a mapping of chromosome numbers on the phylogenetic tree, see Mort et al 2001 Fig. 4.
Subdivision
History
When
While the family can fairly easily be recognised, identifying its constituent genera has been far more problematic.[5] For an extensive history of subfamily Sedoideae, see Ohba 1978. Saint-Hilaire's original description in 1805 included seven genera,[12] as did De Candolle (1815).[13] In a much more extensive treatment in 1828, he divided the Crassulaceae into the two groups, Isostemonae and Diplostemonae (i.e. haplostemony vs. obdiplostemony) on the basis of the number of staminal whorls. The former corresponded to the modern Crassuloideae.[14][5]
Two lineages, six subfamilies, and 33 genera of Crassulaceae were described by Berger in 1930:[15]
Lineages, subfamilies, biogeography, No. genera, type genus (No. species in genus)
- Crassula (Southern hemisphere)
- Crassuloideae S Africa 5 Crassula (300)
- (Crassula, Dinacria, Rochea, Vauanthes, Pagella)
- Kalanchiodeae S Africa, Madagascar 3 Kalanchoe (200)
- (Kalanchöe, Bryophyllum, Kitchingia)
- (
- Cotyledonoideae S Africa, Mediterranean 6 Cotyledon (30)
- (Chiastophyllum, Pistorina, Mucizonia)
- (
- Crassuloideae S Africa 5 Crassula (300)
- Sedum (Northern hemisphere)
- Echeveroideae Mexico 5 Echeveria (150)
- (Echeveria, Villadia, Altamiranoa, Pachyphytum, Lenophyllum)
- Sempervivoideae Mediterranea, Macaronesia 5 Sempervivum (25)
- (Sempervivum, Aeonium, Greenovia, Monanthes, Aichryson)
- Sedoideae N hemisphere, S America, N & E Africa 11 Sedum (500)
- (Sedum (since including Diamorpha), Rosularia, Orostachys, Pseudosedum, Hypagophytum, Afrovivella, Sempervivella, Sinocrassula)
- Echeveroideae Mexico 5 Echeveria (150)
Each of these contained one of the largest genera.
Sedoideae contained three centres of diversity, East Asia, the Mediterranean region and North America, with the greatest in E. Asia. Only a few taxa, such as Rhodiola and Hylotelephium, occurring in all three regions. About 120 species were found in Europe and adjacent parts of North Africa and West Asia,[30] and 400 in Eastern and central Asia.[31]
Within Sedoideae, the large cosmopolitan typical genus Sedum (ca. 500 species), accounts for much of these issues, together with several smaller genera. Sedum refers to herbaceous, predominantly perennial species with alternate and entire leaves, a single subaxial
- subgenus Aizoon
- subgenus Balfouria
- subgenus Spathulata
- subgenus Sedum
- subgenus Telmissa
Grulich (1984) continued this process, proposing
Subsequently, various revisions have proposed fewer subfamilies.
Molecular phylogenetics
Prior to the use of molecular methods of classification, attempts to replace Berger's system were largely unsuccessful.
- Subfamily Crassuloideae Berger Type: Crassula 2 South African genera (250 species)
- Subfamily Sedoideae Berger Type: Sedum
- Tribe Kalanchoeae 't Hart Type: Kalanchoe 5 S African genera (250 spp.)
- Tribe Sedeae 't Hart Type: Sedum
- Subtribe Telephiinae 't Hart Type: Hylotelephium 8 Asian genera (150 spp.)
- Subtribe Sedinae 't Hart Type: Sedum 18 Northern hemisphere genera (700 spp.)
The basal split at subfamily level, separates the
These clades were (1–7):
- Crassula/Crassuloideae, the basal divergence, corresponds to Berger's subfamily of that name and are haplostemonous, but this feature is homoplasious. Confined to southern Africa, except for aquatic species, which are cosmopolitan.[10]
- Kalanchoe/Kalanchoeae, the second divergence, corresponds to Berger's Kalanchoideae (Kalanchoe, Bryophyllum, Kitchingia) and the 2 S. African members of Cotyledonoideae (Adromischus, Cotyledon), together with leaves that are flat, crenate or dentate (toothed), often petiolate and decussate. Chromosome number x=9. Within the clade, Adromischus forms the basal divergence, followed by Cotyledon/Tylecodon as sister to Kalanchoe.[10]
- Telephium/Telephiinae include members of Cotyledonoideae (Umbilicus), together with some Sedoideae genera (including Hylotelephium) and two of Ohba's five subgenera of Sedum (Aizoon and Spathulata)rhizomes (monopodial or sympodial). In this respect many species share leaf features with the Kalanchoeae and Hart considered that the Telephiinae "bridge the gap" between the African Kalanchoeae and the Northern hemisphere Sedinae. Distribution predominantly East Asia, with Umbilicus being Mediterranean.[10]
- Sempervivum includes the montane/alpine Eurasian Sempervivum, its nominative genus, together with Sedum from the same region, including Sedum series Rupestria. Sempervivum is closely related to Jovibarba, which some authors place within the former genus.[10]
- Leucosedum, i.e. "White Sedum", from Sedum album, is polygeneric and includes additional Cotyledonoideae and Eurasian Sedoideae, including Sedum album and other species of Sedum subgenus Gormania. The other genera are thought to have evolved from the Sedum lineage in this clade, including Dudleya and Sedella (N America) and Rosularia, Prometheum and Pistorina (Eurasia). This grouping of 5–7 genera accounts for about 200 species. Some Sedum subgenus Sedum species also place here. Leucosedum species are found throughout the arid southwest United States and Mexico, as well as Eurasia.[9][10]
- Aeonium is predominantly Macaronesian Sempervivoideae (Aeonium, Aichryson, Greenovia and Monanthes), from a N African ancestor, and N. African Sedum. Berger grouped the genera of that subfamily on the basis of polymerous flowers, but this is not restricted to this clade.[9][10]
- Acre, with about 7 genera and 500 species is the most taxon rich clade in the Crasulaceae. It includes the American subfamily Echeveroideae and Sedum from Asia, Europe and Macaronesia, Mexico and Africa, including Sedum acre and Sedum subgenus Sedum.[36] The strong representation of Sedum in this large clade accounts for it comprising a third of the diversity of the family. Two subclades consist of the N American and Macaronesian taxa, the other Eurasian.[9][10]
The last subtribe, the Sedinae, represents the last four clades (4–7) and contained half of the genera and species of Crassulaceae, including Sedum, which is represented in all four clades, and the bulk of clades 5 and 7. In addition to Sedum, 16 other genera are recognised. Aeonium is basal divergence, followed by Sempervivum, with Leucosedum and Acre as sister groups. The Sedinae were very diverse, making phenotypic circumscription impossible. A similar problem exists for each of its subclades.[10][36] Given the realisation that Sedum s.l. was a highly artificial construction, there was support for reducing it by describing a number of segregate genera. Ohba (1995) proposed that Sedum s.s. should be restricted to clade 7, or at most clades 5–7, continuing some of the premolecular work in this direction, newly describing a number of Asian genera in addition to this reduced Sedum.:[31]
- Hylotelephium
- Orostachys
- Aizopsis
- Phedimus
- Rhodiola
- Prometheum
- Rosularia
- Balfouria
- Sinocrassula
- Meterostachys
- Pseudosedum
The general phylogenetic topology described by 't Hart et al. (1995) was confirmed in a larger study of 112 species of Crassulaceae sampled from 33 genera, and all six recognized subfamilies, using the chloroplast gene
Clade1[3] | Hart 1995[2] | Thiede & Eggli 2007[3] | ||||
---|---|---|---|---|---|---|
Subfamily | Tribe | Subtribe | Subfamily | Tribe | ||
1. Crassula | Crassuloideae | Crassuloideae | ||||
2. Kalanchoe | Sedoideae | Kalanchoeae | Kalanchoideae | |||
3. Telephium Hylotelephium2 |
Sedeae | Telephiinae | Sempervivoideae | Telephieae | ||
Rhodiola3 | Umbiliceae | |||||
4. Sempervivum | Sedinae | Semperviveae | ||||
6. Aeonium | Aeonieae | |||||
5. Leucosedum | Sedeae | |||||
7. Acre | ||||||
Notes:
1. Clade numbers following van Ham,[5] order following phylogeny of Thiede & Eggli |
Hart's taxonomic classification was revised by Thiede and Eggli (2007) to define three molecularly defined subfamilies, corresponding to the major
Cladogram of Crassulaceae subfamilies[3] | |||||||||||||||
|
Subfamilies
Crassuloideae Burnett
Crassuloideae is the smallest subfamily, representing a single monophyletic clade (Crassula), defined by
- TillaeaL.) c. 200 spp.
- Hypagophytum A.Berger 1 sp.
Kalanchoöideae A.Berger
Kalanchoöideae is the next smallest subfamily, characterised by flower parts in fours. It represents Berger's Kalanchiodeae and Cotyledonoideae, in part. It is distributed in Madagascar and tropical Africa, with four genera and about between 130–240 species.[38] It is characterised by fused corollas, chromosome number x=9 and mostly southern African distribution. The boundaries between Kalanchoe, Bryophyllum and Kitchingia have remained unclear, and the latter two genera are more commonly treated as sections of Kalanchoe:[3][9][10][37]
- Adromischus Lem.
- Cotyledon L.
- Kalanchoe Adans. (including Bryophyllum Adans., Kitchingia Adans.)
- Tylecodon Toelken
Sempervivoideae Arn.
Sempervivoideae is the largest and taxonomically most complex subfamily, distributed in temperate climates, with about 20–30 genera, and divided into five tribes, of which Sedeae contains two distinct clades, Leucosedum and Acre:[3]
- Telephieae
- Umbiliceae
- Semperviveae
- Aeonieae
- Sedeae
Distribution and habitat
The family Crassulaceae has a cosmopolitan distribution, particularly Crassula, though rare in South America and Australia,
Habitat
Crassulaceae are found predominantly in
Ecology
While most Crassulaceae are perennial, Tillaea are annuals, and annual species are also found among Aichryson, Crassula, Sedum and Monanthes.[11]
Cultivation
Many Crassulaceae species are cultivated as pot plants or in rock gardens and borders.[8]
Toxicity
Some species are toxic to animals, such as those of
Uses
Although no species have a role as
Notes
- ^ Hendrik (Henk) t' Hart 1944–July 2000. Division of Plant Ecophysiology, Utrecht University[6]
References
- ^ a b c d APG IV 2016.
- ^ a b c d e f Hart 1995.
- ^ a b c d e f g h i j k l m Thiede & Eggli 2007.
- ^ a b c d e f Mayuzumi & Ohba 2004.
- ^ a b c d e f g h i j k l m n van Ham & Hart 1998.
- ^ a b c d e Eggli 2003.
- ^ a b c Christenhusz et al 2017.
- ^ a b Hosch 2008.
- ^ a b c d e f g h i j k l m n o p Mort et al 2001.
- ^ a b c d e f g h i j k l m n o p q Gontcharova & Gontcharov 2007.
- ^ a b c d e Mort et al 2010.
- ^ a b Saint-Hilaire 1805.
- ^ a b de Lamarck & de Candolle 1815.
- ^ a b de Candolle 1828.
- ^ a b c d Berger 1930.
- ^ Anonymous 1937.
- ^ a b c d Gontcharova et al 2006.
- ^ a b c d e f g Gontcharova & Gontcharov 2009.
- ^ a b c d e Hart 1997.
- ^ a b c Cronquist 1981.
- ^ Christenhusz & Byng 2016.
- ^ a b Takhtajan 1987.
- ^ a b Thorne 1992.
- ^ a b Stevens 2019.
- ^ Jian et al 2008.
- ^ Linnaeus 1753.
- ^ a b c d Ohba 1978.
- ^ Borisova 1939.
- ^ Borisova 1969.
- ^ a b c Grulich 1984.
- ^ a b c Ohba 1995.
- ^ Fröderströmm 1935.
- ^ Carillo-Reyes et al 2009.
- ^ Nikulin, Nikulin & Gontcharov 2015.
- ^ Tölken 1978.
- ^ a b Hart 1995a.
- ^ a b Rosen 2018.
- ^ a b Messerschmid et al 2020.
- ^ a b Wiersema & León 2016.
- ^ Gwaltney-Brant 2012.
Bibliography
Books
- Borisova, AG (1939). "Crassulaceae D.C.". In Komarov, VL (ed.). Flora of the U.S.S.R. IX Rosales and Sarraceniales. Translated by Israel Program for Scientific Translations. Moskva: Botanicheskii institut Akademiia Nauk SSSR [Botanical Institute of the Academy of Sciences USSR]. pp. 8–105, addenda 357–372.
- ISBN 978-0-226-52292-0.
- ISBN 978-0-231-03880-5.
- Eggli, Urs; Newton, Leonard E. (2004). Etymological Dictionary of Succulent Plant Names. ISBN 978-3-540-00489-9.
- Eggli, Urs, ed. (2003). Illustrated Handbook of Succulent Plants: Crassulaceae. S2CID 36280482.
- Fröderströmm, Harald (1935). The Genus Sedum L.: a systematic essay. Meddelanden från Göteborgs botaniska trädgärd vol. 7. Elander Boktryckeri Aktiebolag.
- Hart, H. 't; Eggli, U., eds. (1995). Evolution and systematics of the Crassulaceae (23rd Congress of the International Organization for Succulent Plant Study, Wageningen, Netherlands, August 20th, 1994). Leiden: Backhuys. OCLC 34335028.
- Mitchell, John; ISBN 978-0-88978-047-7.
- Peterson, Michael E.; Talcott, Patricia A. (2012). Small Animal Toxicology. ISBN 978-1-4557-0717-1.
- Smith, Gideon F.; Figueiredo, Estrela; Wyk, Abraham E. van (2019). Kalanchoe (Crassulaceae) in Southern Africa: Classification, Biology, and Cultivation. ISBN 978-0-12-814008-6.
- Takhtajan, A. (1987). Sistema Magnoliofitov (Systema Magnoliophytorum) (in Russian). Leningrad: Nauka.
- Thiede, J; Eggli, U (2007). "Crassulaceae". In )
- Wiersema, John H.; León, Blanca (2016) [1999]. World Economic Plants: A Standard Reference (2nd ed.). ISBN 978-1-4665-7681-0.
Chapters
- Gwaltney-Brant, Sharon M. (2012). Christmastime Plants. pp. 419–512., in Peterson & Talcott (2012)
- Hart, H. 't; Eggli, U. (1995). Introduction. pp. 7–15., in Hart & Eggli (1995)
- Hart, H. 't (1995). Infrafamilial and generic classification of the Crassulaceae. pp. 159–172., in Hart & Eggli (1995)
- Ohba, Hideaki (1995). Systematic problems of Asian Sedoideae. pp. 151–158., in Hart & Eggli (1995)
- van Ham, R.C.H.J. (1995). Phylogenetic relationships in the Crassulaceae inferred from chloroplast DNA variation. pp. 16–29., in Hart & Eggli (1995)
Historical
- Berger, A. (1930). "Crassulacaeae". In Engler, Adolf; Prantl, Karl Anton (eds.). Die Natürlichen Pflanzenfamilien. Vol. 18A. Leipzig: Verlag von Wilhelm Engelmann. pp. 352–483.
- Borisova, AG (1969). "Conspectus Systematis Fam. Crassulaceae D.C. Florae URSS". Novit. Syst. Pl. Vasc. Acad. Sci. USSR. 6: 112–121.
- de Candolle, A. P. (1828). "Crassulaceae". Prodromus systematis naturalis regni vegetabilis, sive, Enumeratio contracta ordinum generum specierumque plantarum huc usque cognitarium, juxta methodi naturalis, normas digesta. Vol. 3. Paris: Treuttel et Würtz. pp. 381–414.
- . Vol. 4 (3 ed.). Paris: Desray. pp. 382–398.
- Gray, Samuel Frederick (1821). "Crassulaceae". A natural arrangement of British plants: according to their relations to each other as pointed out by Jussieu, De Candolle, Brown, &c. 2 vols. London: Baldwin, Cradock, and Joy. pp. ii: 538–543.
- Linnaeus, Carl (1753). Species Plantarum: exhibentes plantas rite cognitas, ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas. Stockholm: Impensis Laurentii Salvii. pp. 282–283, 128–129, 429–30, 464–465, 1035, 430–432., see also Species Plantarum
- Saint-Hilaire, Jean Henri Jaume (1805). "Crassuleae". Exposition des familles naturelles et de la germination des plantes, contentant la description de 2,337 genres et d'environ 4,000 espèces, 112 planches dont les figures ont été dessinées par l'auteur. vol II. Paris: Treuttel et Würtz. pp. 123–128.
Articles
- Anonymous (1937). "The Crassulaceae". The Cactus Journal. 5 (3): 51–53. JSTOR 42784672.
- .
- Grulich, Vit (1984). "Generic division of Sedoideae in Europe and the adjacent regions" (PDF). Preslia. 56: 29–45.
- Hart, H. 't; Koek-Noorman, J. (November 1989). "The origin of the woody Sedoideae (Crassulaceae)". JSTOR 1222628.
- Ohba, Hideaki (March 1977). "The taxonomic status of Sedum telephium and its allied species (Crassulaceae)". The Botanical Magazine Tokyo. 90 (1): 41–56. S2CID 22239507.
- Ohba, H (1978). "Generic and infrageneric classification of the old world sedoideae crassulaceae". Journal of the Faculty of Science University of Tokyo Section III Botany. 12 (4): 139–193.
- S2CID 40348158.
Phylogeny
- Angiosperm Phylogeny Group (2016). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV". .
- Carrillo-Reyes, Pablo; Sosa, Victoria; Mort, Mark E. (October 2009). "Molecular phylogeny of the Acre clade (Crassulaceae): Dealing with the lack of definitions for Echeveria and Sedum" (PDF). PMID 19482091. Archived from the original(PDF) on 2019-08-24. Retrieved 2019-08-24.
- Gontcharova, S. B.; Artyukova, E. V.; Gontcharov, A. A. (June 2006). "Phylogenetic relationships among members of the subfamily Sedoideae (Crassulaceae) inferred from the ITS region sequences of nuclear rDNA" (PDF). Russian Journal of Genetics. 42 (6): 654–661. S2CID 33249840. Archived from the original(PDF) on 2017-08-09. Retrieved 2019-09-27.
- Gontcharova, Svetlana B.; Gontcharov, Andrey A. (2007). "Molecular Phylogenetics of Crassulaceae" (PDF). Genes, Genomes and Genomics. 1 (1): 40–46.
- Gontcharova, S. B.; Gontcharov, A. A. (11 October 2009). "Molecular phylogeny and systematics of flowering plants of the family Crassulaceae DC". S2CID 39602334.
- Hart, H. 't (1995a). "The evolution of the Sedum acre group (Crassulaceae)" (PDF). Bocconea. 5: 119–128.
- Hart, H.'t (1997). "Diversity within Mediterranean Crassulaceae" (PDF). Lagascalia. 1 (2): 93–100.
- Jian, Shuguang; PMID 18275001.
- Mayuzumi, Shinzo; Ohba, Hideaki (2004). "The Phylogenetic Position of Eastern Asian Sedoideae (Crassulaceae) Inferred from Chloroplast and Nuclear DNA Sequences". S2CID 84319808.
- Messerschmid, Thibaud F.E.; Klein, Johannes T.; Kadereit, Gudrun; Kadereit, Joachim W. (4 September 2020). "Linnaeus's folly – phylogeny, evolution and classification of Sedum (Crassulaceae) and Crassulaceae subfamily Sempervivoideae". .
- Mort, Mark E.; PMID 11159129.
- Mort, Mark E.; Levsen, Nicholas; Randle, Christopher P.; Van Jaarsveld, Ernst; Palmer, Annie (July 2005). "Phylogenetics and diversification of Cotyledon (Crassulaceae) inferred from nuclear and chloroplast DNA sequence data". PMID 21646139.
- Mort, Mark E.; O'Leary, T. Ryan; Carrillo-Reyes, Pablo; Nowell, Tracey; Archibald, Jenny K.; Randle, Christopher P. (December 2010). "Phylogeny and evolution of Crassulaceae: Past, present, and future". Biodiversity & Ecology. 3: 69–86.
- Nikulin, Arthur; Nikulin, Vyacheslav; Gontcharov, Andrey (October 2015). "To the question of phylogenetic structure of the tribe Telephieae (Sempervivoideae, Crassulaceae) based on ITS rDNA sequence comparisons". Botanicheskii Zhurnal (in Russian). 100 (10): 1030–1040.
- Nikulin, Vyacheslav Yu.; Gontcharova, Svetlana B.; Stephenson, Ray; Gontcharov, Andrey A. (September 2016). "Phylogenetic relationships between Sedum L. and related genera (Crassulaceae) based on ITS rDNA sequence comparisons". Flora. 224: 218–229. .
- van Ham, Roeland C. H. J.; Hart, Henk ’t (January 1998). "Phylogenetic relationships in the Crassulaceae inferred from chloroplast DNA restriction-site variation". PMID 21684886.
- PMID 23629845.
- Tölken, H. (15 December 1978). "New taxa and new combinations in Cotyledon and allied genera". .
Websites
- Hosch, William L. (27 Jun 2008). Crassulaceae. Encyclopædia Britannica. Retrieved 18 August 2019.
- Stevens, P.F. (2019) [2001]. "Crassulaceae". AP Web v. 14. Missouri Botanical Garden. Retrieved 31 July 2019. (see also Angiosperm Phylogeny Website)
- Fu, Kunjun; Ohba, Hideaki; Gilbert, Michael G. (2004). "Crassulaceae Candolle". p. 202. Retrieved 24 August 2019., in Flora of China online vol. 8
- Rosen, Ralph (5 April 2018). "Crassuloideae". The Cactus Expert: World of Succulents. Retrieved 18 August 2019.
- "International Crassulaceae Network". Retrieved 5 September 2019.
- Etter, Julia; Kristen, Martin (2019). "The Crassulaceae database". Retrieved 9 October 2019.
Databases
- "The Plant List Version 1.1: Crassulaceae". The Plant List. Royal Botanic Gardens, Kew and Missouri Botanical Garden. 2013. Retrieved 1 September 2019.
- "Crassulaceae J.St.-Hil". World Flora Online. 2019. Retrieved 1 September 2019.
- "Crassulaceae J.St.-Hil". Plants of the World Online. Royal Botanic Gardens, Kew. 2019. Retrieved 1 September 2019.
External links
- Media related to Crassulaceae at Wikimedia Commons
- Data related to Crassulaceae at Wikispecies