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Ranunculaceae

{{Short description|Family of eudicot flowering plants}}

{{Automatic taxobox

| fossil_range = {{fossil range|124|0}} Early Cretaceous–Recent{{sfn|Pigg|DeVore|2005}}

| image = Ranunculus_auricomis.jpg

| image_caption = Ranunculus auricomus

| taxon = Ranunculaceae

| authority = Juss.

| type_genus = Ranunculus

| type_genus_authority = L.

| subdivision_ranks = Subfamilies

| subdivision = *Hydrastidoideae

}}

Ranunculaceae ({{IPAc-en|r|@|n|V|ng|k|j|u:|'|l|eI|s|i|%|aI|,_|-|s|i:|%|i:}}, buttercup or crowfoot family; Latin {{lang|la|rānunculus}} "little frog", from {{lang|la|rāna}} "frog") is a family of over 2,000 known species of flowering plants in 43 genera,{{sfn|Christenhusz|Byng| 2016}} distributed worldwide.

The largest genera are Ranunculus (600 species), Delphinium (365), Thalictrum (330), Clematis (380), and Aconitum (300).

Description

File:Adonis autumnalis flowerdiagram.png. Adonis annua]]

Ranunculaceae are mostly herbaceous annuals or perennials, but some are woody climbers (such as Clematis){{sfn|Clapham|Tutin|Warburg|1981}} or shrubs (e.g. Xanthorhiza).

Most members of the family have bisexual flowers which can be showy or inconspicuous. Flowers are solitary, but are also found aggregated in cymes, panicles, or spikes. The flowers are usually radially symmetrical but are also found to be bilaterally symmetrical in the genera Aconitum and Delphinium.{{sfn|Stace|2010}}{{sfn|Ronse de Craene|2010}} The sepals, petals, stamens and carpels are all generally free (not fused), the outer flower segments typically number four or five. The outer stamens{{efn|In the Ranunculaceae, a variety of terms are used to describe the whorl of structures between the sepals and stamens, including honey-leaves, petals, staminodes or nectaries{{sfn|Tamura|1993|p=564}}}} may be modified to produce only nectar, as in Aquilegia, Helleborus and Delphinium.{{sfn|Ronse de Craene|2010}}

In some genera, such as Thalictrum, the sepals are colorful and appear petal-like (petaloid) and the petals can be inconspicuous or absent.{{sfn|Clapham|Tutin|Warburg|1981}} The stems are unarmed. The leaves are variable. Most species have both basal and cauline (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 rhizomes that develop new roots each year. Ficaria verna can reproduce vegetatively by means of root tubers produced in the leaf axils.{{sfn|Clapham|Tutin|Warburg|1981}}{{sfn|Stace|2010}} Some members of the genus Thalictrum utilize anemophily while others utilize entomophily.{{cite journal |last1=Steven |first1=Janet |last2=Waller |first2=Donald |date=2004 |title=Reproductive alternatives to insect pollination in four species of Thalictrum (Ranunculaceae) |url=https://www.researchgate.net/publication/227512644 |journal=Plant Species Biology |volume=19 |issue=2 |pages=73–80 |doi=10.1111/j.1442-1984.2004.00103.x |access-date=6 December 2018 }} Flowers of the entomophilous genus Papaver, also of the Ranunculales order, produce only pollen.{{cite web |url=https://www.beeculture.com/treat-bees-banquet-poppies/ |title=Treat Your Bees to a Banquet of Poppies |last=Ross |first=Gary |publisher=A.I. Root Company |date=22 December 2016 |website=Bee Culture |access-date=6 December 2018}} Until recently, it was believed that the species of the genus Anemone also lack nectar.{{cite journal |last1=Erbar |first1=Claudia |last2=Leins |first2=Peter |date=2013 |title=Nectar production in the pollen flower of Anemone nemorosa in comparison with other Ranunculaceae and Magnolia (Magnoliaceae) |url=https://www.researchgate.net/publication/257796107 |journal=Organisms Diversity & Evolution |volume=13 |issue=3 |pages=287–300 |doi=10.1007/s13127-013-0131-9 |bibcode=2013ODivE..13..287E |s2cid=16275166 |access-date=6 December 2018 }}

The fruits are most commonly free, unfused achenes (e.g. Ranunculus, Clematis) or follicles (e.g. Helleborus, Eranthis, Nigella), but a berry in Actaea.{{sfn|Clapham|Tutin|Warburg|1981}}{{sfn|Stace|2010}}

{{multiple image

| perrow = 2

| header = Fruit Morphology

| align = right

| image1 = Pulsatilla alpina fruit.JPG

| caption1 = Achene: Pulsatilla alpina

| image2 = Ranunculus acris0.jpg

| caption2 = Achene: Ranunculus acris

| image4 = Helleborus niger (Früchte).jpg

| caption4 = Follicle: Helleborus niger

| image3 = Nigella_arvensis_fruit.jpg

| caption3 = Follicle: Nigella arvensis

}}

= Phytochemistry =

Ranunculaceae 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.{{Cite web|title=Anemone (Windflower) {{!}} North Carolina Extension Gardener Plant Toolbox|url=https://plants.ces.ncsu.edu/plants/anemone/|access-date=2021-02-17|website=plants.ces.ncsu.edu}}

Other poisonous or toxic compounds, alkaloids and glycosides, are also common.{{citation needed|date=April 2018}}

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.[http://botany.csdl.tamu.edu/FLORA/newgate/cronang.htm Flowering Plant Gateway] David, (2010){{cite web|url = http://apps.rhs.org.uk/rhsplantfinder/documents/Essay.pdf |title = Plants in their proper places – the new classification of flowering plants|publisher = Royal Horticultural Society|archive-date = 18 July 2011 |archive-url = https://web.archive.org/web/20110718073537/http://apps.rhs.org.uk/rhsplantfinder/documents/Essay.pdf}} 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.{{sfn|APG|2016}} The family is monophyletic with Glaucidium as sister to the remaining genera.{{sfn|Wang et al|2009}} This phylogeny is illustrated in the APG Poster.[https://www.bcp.fu-berlin.de/biologie/arbeitsgruppen/Archiv/ag_hilger/publikationen/poster1.pdf 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 Paeonia, 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 Paeonia and Glaucidium were no longer considered to belong to Ranunculaceae.{{sfn|Tamura|1993}} 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

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.{{sfn|Stevens|2017}}{{sfn|Wang et al|2009}} Both genera are represented by a single species, Glaucidium palmatum and Hydrastis canadense 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.{{sfn|Wang et al|2009}} 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 (see below).

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.{{Cite journal |last1=He |first1=Jian |last2=Lyu |first2=Rudan |last3=Luo |first3=Yike |last4=Xiao |first4=Jiamin |last5=Xie |first5=Lei |last6=Wen |first6=Jun |last7=Li |first7=Wenhe |last8=Pei |first8=Linying |last9=Cheng |first9=Jin |date=September 2022 |title=A phylotranscriptome study using silica gel-dried leaf tissues produces an updated robust phylogeny of Ranunculaceae |url=https://linkinghub.elsevier.com/retrieve/pii/S1055790322001580 |journal=Molecular Phylogenetics and Evolution |language=en |volume=174 |pages=107545 |doi=10.1016/j.ympev.2022.107545|pmid=35690374 |bibcode=2022MolPE.17407545H }}

;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.

{{cladogram|title=Cladogram of Ranunculaceae subfamilies{{sfn|Wang et al|2009}}

|align=center

|cladogram={{clade|style=width:600px;

|label1= Ranunculaceae

|1={{clade

|1= Glaucidoideae80px

|2={{clade

| 1=Hydrastidoideae80px

| 2={{clade

| 1=Coptidoideae80px

| 2={{clade|1=Ranunculoideae80px

| 2=Thalictroideae80px

}}

}}

}}

}}}}}}

{{clear}}

= Genera =

Ranunculaceae contains approximately 43 genera.{{sfn|Christenhusz|Byng| 2016}}

{{div col|colwidth=22em}}

; Subfamily Glaucidioideae

; 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

{{div col end}}

Previous genera

  • Anemonella Spach → Thalictrum
  • Psychrophila (DC.) Bercht. & J.Presl → Caltha

= Fossil record =

Contrary to earlier data, there are no confirmed pre-Tertiary ranunculacean fossils.{{Cite book |last1=Friis |first1=Else Marie |title=Early flowers and angiosperm evolution |last2=Crane |first2=Peter R. |last3=Pedersen |first3=Kaj Raunsgaard |last4=Knorring |first4=Pollyanna von |date=2011 |publisher=Cambridge Univ. Press |isbn=978-0-521-59283-3 |edition=1. publ |location=Cambridge}} The Early Cretaceous Leefructus mirus from the Yixian Formation in China was described as belonging to the "compare Family stem lineage to the Ranunculaceae".{{Cite journal |last1=Sun |first1=Ge |last2=Dilcher |first2=David L. |last3=Wang |first3=Hongshang |last4=Chen |first4=Zhiduan |title=A eudicot from the Early Cretaceous of China |url=https://www.researchgate.net/publication/50940368 |journal=Nature |date=2011 |volume=471 |issue=7340 |pages=625–628|doi=10.1038/nature09811 |pmid=21455178 |bibcode=2011Natur.471..625S }} The oldest confirmed representative of the family is Paleoactaea nagelii Pigg & DeVore described on the basis of fruits coming from the Upper Paleocene of North Dakota.{{sfn|Pigg|DeVore|2005}}

Uses

Some Ranunculaceae are used as herbal medicines because of their alkaloids and glycosides, such as Hydrastis canadensis (goldenseal), whose root is used as a tonic. More than 30 species are used in homeopathy, including Aconitum napellus, Actaea racemosa, Clematis recta, Clematis virginiana, Hydrastis canadensis, Ranunculus bulbosus, Helleborus niger, Delphinium staphisagria, Pulsatilla nigricans. Many genera are well known as cultivated flowers, such as Aconitum (monkshood), Clematis, Consolida (larkspur), Delphinium, Helleborus (Christmas rose), Trollius (globeflower). The seeds of Nigella sativa are used as a spice in Indian and Middle Eastern cuisine.{{sfn|Turner|1984}}

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 hondoense

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

{{Notelist}}

References

{{sfn whitelist|CITEREFFNA}}

{{Reflist|20em|refs=

{{harvnb|FNA|}}

{{harvnb|The Plant List|2013|loc=[http://www.theplantlist.org/1.1/browse/A/Ranunculaceae/ Ranunculaceae]}}

}}

Bibliography

{{Refbegin|30em}}

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  • {{cite book|editor-last1=Kubitzki|editor-first1= Klaus|editor-last2=Rohwer|editor-first2=Jens G.|editor-last3=Bittrich|editor-first3=Volker|title=The Families and Genera of Vascular Plants. II Flowering plants - Dicotyledons. Magnoliid, Hamamelid and Caryophyllid families|url=https://books.google.com/books?id=yjlzrzbRXNQC|date=1993|publisher=Springer|location=Berlin|isbn=978-3540555094|ref={{harvid|Kubitzki et al|1993}}}}
  • {{cite book|last1=Tamura|first1=M|author-link=Minoru N. Tamura|title=Ranunculaceae|pages=563–583|url=https://books.google.com/books?id=yjlzrzbRXNQC&pg=PA563|date=1993-07-28|publisher=Springer|isbn=9783540555094}}, in {{harvtxt|Kubitzki et al|1993}}
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{{Refend}}

External links

{{Commons category}}

{{Wikispecies|Ranunculaceae}}

{{Refbegin|2}}

  • [http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=10757 Flora of North America: Ranunculaceae]
  • [http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=10757 Flora of China: Ranunculaceae]
  • [https://web.archive.org/web/20080524060601/http://www.topwalks.net/plants/generos/ranunculaceae_01.htm Ranunculaceae in Topwalks]
  • [https://www.botanical-dermatology-database.info/BotDermFolder/RANU.html Ranunculaceae] in [https://www.botanical-dermatology-database.info/index.html BoDD – Botanical Dermatology Database]
  • [https://www.delta-intkey.com/angio/www/ranuncul.htm Ranunculaceae] in L. Watson and M.J. Dallwitz (1992 onwards). [http://delta-intkey.com/angio/ The families of flowering plants].
  • [https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=3440&lvl=3&p=mapview&p=has_linkout&p=blast_url&p=genome_blast&lin=f&keep=1&srchmode=1&unlock NCBI Taxonomy Browser]
  • [https://web.archive.org/web/19990824065228/http://www.csdl.tamu.edu/FLORA/cgi/gateway_family?fam=ranunculaceae Links at CSDL, Texas]
  • [https://web.archive.org/web/20070930184616/http://www.alpine-plants-jp.com/art/index_kinpougeka.htm Japanese Ranunculaceae] - Flavon's art gallery
  • [http://www.ajol.info/index.php/ajb/article/viewFile/65953/53657 Cai et al. Molecular phylogeny of Ranunculaceae based on internal transcribed spacer sequences 2009]
  • [https://indianmedicinalplants.info/d2/index.htm Aconitum heterophyllum (fam. Ranunculaceae) at the Encyclopedia of Ayurvedic Medicinal Plants]

{{Refend}}

{{Taxonbar|from=Q145869}}

{{Authority control}}

Category:Eudicot families

Category:Extant Cretaceous first appearances