In some genera, such asThalictrum, the sepals are colorful and appear petal-like (petaloid) and the petals can be inconspicuous or absent.[3] The stems are unarmed. The leaves are variable. Most species have bothbasal andcauline (stem) leaves, which are usually compound or lobed but can be simple. They are typically alternate, or occasionally opposite or even whorled. Many species, especially the perennials, formrhizomes that develop new roots each year.[6]Ficaria verna can reproduce vegetatively by means of root tubers produced in the leaf axils.[3][4] Some members of the genusThalictrum utilizeanemophily while others utilizeentomophily.[8] Flowers of the entomophilous genusPapaver, also of theRanunculales order, produce onlypollen.[9] Until recently, it was believed that the species of the genusAnemone also lacknectar.[10]
Many Ranunculaceae members containprotoanemonin, which is toxic to humans and animals. Contact with plant sap may cause inflammation and blistering of the skin, while ingestion can cause irritation of the mouth, vomiting, and diarrhea.[11]Toxic alkaloids such asaconitine may also be present in some species such asAconitum carmichaelii.[citation needed]
Takhtajan (1997) included the Ranunculaceae as the only family in the Ranunculales which he placed in a subclass, the Ranunculidae, instead of a superorder. Previously, Thorn (1992) placed the Ranunculaceae in the Berberidales, an order within the Superorder Magnolianae. Earlier Cronquist in 1981 included the Ranunculaceae along with seven other families in the Rancunculales which was included in the Magnoliidae, which he regarded as a subclass.[12] David, (2010)[13] placed the Ranuculaceae, together with theEupteleaceae,Lardizabalaceae,Menispermaceae,Berberidaceae, andPapaveraceae in the Ranunculales, the only order in the superorder Ranunculanae. This follows the work of theAngiosperm Phylogeny Group.[citation needed]
Early subdivisions of the family, such asMichel Adanson (1763), simply divided it based on one-seeded or many-seeded fruit.Prantl (1887) envisaged threetribes, Paeonieae, Helleboreae and Anemoneae withPaeonia,Glaucidium andHydrastis forming Paeonieae. By the twentieth century Langlet (1932) used chromosome types to create twosubfamilies, Ranunculoideae and Thalictroideae. In 1966,Tamura further developed Langlet's system by adding floral characteristics with six subfamilies;[citation needed]
but by 1988 he had reduced Coptidoideae to a tribe within Isopyroideae, leaving five subfamilies, an arrangement he continued in his 1993 monograph, dividing the larger subfamilies into tribes, though by thenPaeonia andGlaucidium were no longer considered to belong to Ranunculaceae.[17]Paeonia was separated from Ranuculaceae and placed in its own family ofPaeoniaceae (orderSaxifragales). Other genera originally included in Ranunculaceae includeCircaeaster, which was placed in its own familyCircaeasteraceae.[citation needed]
Tamura's complete system was structured as follows;
When subjected tomolecular phylogenetic analysis, only Thalictroideae is monophyletic. The position ofGlaucidium and some of its unique morphological characteristics prompted Stevens to suggest that it be given subfamilial rank as the monotypic Glaucidioideae. Similarly,Hydrastis has been assigned to subfamily Hydrastidoideae.[18][15] Both genera are represented by a single species,Glaucidium palmatum andHydrastis canadense respectively.[citation needed]
The relationships between the genera suggest the existence of three majorclades corresponding to Coptidoideae, Thalictroideae (clade A) and Ranunculoideae (clade F). The latter is the largest with four subclades (B–E). Of these, C corresponds to Delphineae, D to Cimicifugae and E to Ranunculoideae.[15] Consequently, Wang and colleagues (2009) proposed a new classification with five subfamilies, and further subdividing Ranunculoideae into ten tribes. The relationship between the subfamilies is shown in thecladogram[citation needed]
In addition to the two monotypic subgenera, Coptoideae has 17 species and Thalictroideae has 450, includingThalictrum andAquilegia. The other genera (2025 species, 81% of the family) belong to Ranunculoideae.Kingdonia had been included by Tamura in Anemoneae, but is now added to Circaeasteraceae.[citation needed]
In recent years, researchers have used nuclear genes (obtained through transcriptome sequencing technology) to further investigate the phylogenetic relationships of the Ranunculaceae family, and their findings are consistent with those of Wang and colleagues (2009). However, this study indicates that the ancestors of the Ranunculaceae family experienced multiple whole-genome duplication (WGD) events, which may be related to the longstanding prosperity of this ancient group.[19]
Subfamilies of Ranunculaceae (5) and tribes of Ranunculoideae
Contrary to earlier data, there are no confirmed pre-Tertiary ranunculacean fossils.[21] The Early CretaceousLeefructus mirus from theYixian Formation in China was described as belonging to the "compare Family stem lineage to the Ranunculaceae".[22] The oldest confirmed representative of the family isPaleoactaea nagelii Pigg & DeVore described on the basis of fruits coming from the UpperPaleocene ofNorth Dakota.[1]
^In the Ranunculaceae, a variety of terms are used to describe the whorl of structures between thesepals and stamens, including honey-leaves, petals, staminodes ornectaries[6][7]
Clapham, A.R.; Tutin, T.G.; Warburg, E.F. (1981).Excursion flora of the British Isles (3 ed.). Cambridge, U.K.: Cambridge University Press.ISBN978-0-521-23290-6.: 25
Ronse de Craene, L.P. (2010).Floral diagrams, an aid to understanding flower morphology and evolution. Cambridge University Press.ISBN978-0-521-72945-1.: 140
Wang, Wei; Lu, An-Ming; Ren, Yi; Endress, Mary E.; Chen, Zhi-Duan (January 2009). "Phylogeny and classification of Ranunculales: Evidence from four molecular loci and morphological data".Perspectives in Plant Ecology, Evolution and Systematics.11 (2):81–110.Bibcode:2009PPEES..11...81W.doi:10.1016/j.ppees.2009.01.001.
Turner, NJ (July 1984). "Counter-irritant and other medicinal uses of plants in Ranunculaceae by native peoples in British Columbia and neighbouring areas".Journal of Ethnopharmacology.11 (2):181–201.doi:10.1016/0378-8741(84)90038-2.PMID6387285.
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