Ranunculaceae ( , buttercup or crowfoot family; Latin rānunculus "little frog", from rāna "frog") is a family of over 2,000 known species of in 43 genera, distributed worldwide.
The largest genera are Ranunculus (600 species), Delphinium (365), Thalictrum (330), Clematis (380), and Aconitum (300).
Description
Ranunculaceae are mostly
herbaceous annuals or perennials, but some are woody climbers (such as
Clematis) or shrubs (e.g.
Xanthorhiza).
Most members of the family have which can be showy or inconspicuous. Flowers are solitary, but are also found aggregated in cymes, , or spikes. The flowers are usually radially symmetrical but are also found to be bilaterally symmetrical in the genera Aconitum and Delphinium. The , , and are all generally free (not fused), the outer flower segments typically number four or five. The outer stamens may be modified to produce only nectar, as in Aquilegia, Helleborus and Delphinium.
In some genera, such as Thalictrum, the sepals are colorful and appear petal-like (petaloid) and the petals can be inconspicuous or absent. The stems are unarmed. The leaves are variable. Most species have both basal leaf and cauline leaf (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, form that develop new roots each year.
The are most commonly free, unfused (e.g. Ranunculus, Clematis) or follicles (e.g. Helleborus, Eranthis, Nigella), but a berry in Actaea.
Phytochemistry
Many Ranunculaceae members contain
protoanemonin, 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.
Toxic alkaloids such as
aconitine may also be present in some species such as
Aconitum carmichaelii.
Taxonomy
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.
David, (2010)
placed the Ranuculaceae, together with the
Eupteleaceae,
Lardizabalaceae,
Menispermaceae,
Berberidaceae, and
Papaveraceae in the Ranunculales, the only order in the superorder Ranunculanae. This follows the work of the Angiosperm Phylogeny Group.
The family Ranunculaceae sensu stricto is one of seven families included in the order Ranunculales within the eudicots according to the Angiosperm Phylogeny Group (APG) classification. The family is monophyletic with Glaucidium as sister group to the remaining genera. This phylogeny is illustrated in the APG Poster.[ Angiosperm Phylogeny poster]
Subdivision
Early subdivisions of the family, such as
Michel Adanson (1763), simply divided it based on one-seeded or many-seeded fruit. Prantl (1887) envisaged three tribes, Paeonieae, Helleboreae and Anemoneae with
Peony,
Glaucidium and
Hydrastis forming Paeonieae. By the twentieth century Langlet (1932) used chromosome types to create two
subfamilies, Ranunculoideae and Thalictroideae. In 1966, Tamura further developed Langlet's system by adding floral characteristics with six subfamilies;
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 then Peony and Glaucidium were no longer considered to belong to Ranunculaceae. Paeonia was separated from Ranuculaceae and placed in its own family of Paeoniaceae (order Saxifragales). Other genera originally included in Ranunculaceae include Circaeaster, which was placed in its own family Circaeasteraceae.
Tamura's complete system was structured as follows;
- Subfamilies and tribes
-
Subfamily Ranunculoideae Hutch.
-
Subfamily Helleboroideae Hutch.
-
Subfamily Isopyroideae Tamura
-
Subfamily Thalictroideae
-
Subfamily Hydrastidoideae
The genus Glaucidium, having been moved to its own family (Glaucidiaceae), has since been restored to Ranuculaceae.
Molecular phylogenetics
When subjected to molecular phylogenetic analysis, only Thalictroideae is monophyletic. The position of
Glaucidium 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. Both genera are represented by a single species,
Glaucidium palmatum and
Hydrastis respectively.
The relationships between the genera suggest the existence of three major clades 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. 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 the cladogram
In addition to the two monotypic subgenera, Coptoideae has 17 species and Thalictroideae has 450, including Thalictrum and Aquilegia. 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.
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.
- Subfamilies of Ranunculaceae (5) and tribes of Ranunculoideae
-
Glaucidioideae (Tamura) Loconte (1)
-
Hydrastidoideae Engler (1)
-
Coptidoideae Tamura (2)
-
Thalictroideae Raf. (10)
-
Ranunculoideae Arn. (46)
-
Adonideae Kunth
-
Delphinieae Schröd.
-
Nigelleae Schröd.
-
Helleboreae DC.
-
Cimicifugeae Torr. and A.Gray
-
Caltheae Bercht. and J.Presl
-
Asteropyreae W.T.Wang and C. Y.Chang
-
Callianthemeae W.Wang and Z. D.Chen
-
Anemoneae DC.
-
Ranunculeae DC.
Genera
Ranunculaceae contains approximately 43 genera.
[
]
- Subfamily Glaucidioideae
-
Glaucidium Siebold & Zuccarini
- Subfamily Hydrastidoideae
- Subfamily Coptidoideae
- Subfamily Thalictroideae
- Subfamily Ranunculoideae
- ; Tribe Adonideae
- ; Tribe Delphinieae
- ; Tribe Nigelleae
- ; Tribe Helleboreae
- ; Tribe Cimicifugeae
- ; Tribe Caltheae
- ; Tribe Asteropyreae
- ; Tribe Callianthemeae
- ; Tribe Anemoneae
- ; Tribe Ranunculeae
Previous genera
-
Anemonella Spach → Thalictrum
-
Psychrophila (DC.) Bercht. & J.Presl → Caltha
Fossil record
Contrary to earlier data, there are no confirmed pre-Tertiary period ranunculacean fossils.
Uses
More than 30 species of Ranunculaceae are used in traditional medicines, including Aconitum napellus, Actaea racemosa, Clematis recta, Clematis virginiana, Hydrastis canadensis, Ranunculus bulbosus, Helleborus niger, Delphinium staphisagria, Pulsatilla nigricans. Many genera are commonly grown in , such as Aconitum (monkshood), Clematis, Aquilegia, Consolida (larkspur), Delphinium, Helleborus (Christmas rose), Trollius (globeflower). The seeds of Nigella sativa are used as a spice in Indian and Middle Eastern cuisine.
Gallery
Tribes of subfamily Ranunculoideae
Image:Adonis_annua_flor.jpg|Adonideae:
Adonis annua
Image:Delphinium_elatum_hybride_001.jpg|Delphinieae:
Delphinium elatum hybrid
Image:2007-10-25Nigella_damascena_09.jpg|Nigelleae:
Nigella damascena
Image:Lenteroos frekkels.jpg|Helleboreae:
Helleborus × hybridus
Image:Actaea simplex and Parnara guttata in Mount Ibuki 2011-08-28.jpg|Cimicifugeae:
Actaea simplex
Image:2007-03-27Caltha_palustris02.jpg|Caltheae:
Caltha palustris
Image:Callianthemum hondoense 01.jpg|Callianthemeae:
Callianthemum
Image:10_kwietnia_2009_r._Zawilec_gajowy_033.jpg|Anemoneae:
Anemone nemorosa
Image:Ranunculus auricomis.jpg|Ranunculeae:
Ranunculus auricomus
Other subfamilies
File:Coptis trifolia 5.JPG|Coptidoideae:
Coptis trifolia
File:Glaucidium palmatum 0a4.JPG|Glaucidioideae:
Glaucidium palmatum
File:Hydrastis vision general.jpg|Hydrastidoideae:
Hydrastis canadensis
File:Thalictrum delavayi Prague 2013 4.jpg|Thalictroideae:
Thalictrum delavayi
File:Akleja.Aquilegia vulgaris.jpg|Thalictroideae:
Aquilegia vulgaris
File:Isopyrum thalictroides2.jpg|Thalictroideae:
Isopyrum thalictroides
Notes
Bibliography
-
-
-
-
Langlet, O. (1932). Uber Chromosomenverhaltnisse und Systematik der Ranunculaceae. Svensk Bot. Tidskr 26, 381–401.
-
-
Strasburger, Noll, Schenck, Schimper: Lehrbuch der Botanik für Hochschulen. 4. Auflage, Gustav Fischer, Jena 1900, p. 459 (flower diagrams)
-
Sandro Pignatti. Flora d'Italia, Edagricole, Bologna 1982.
External links
