Papaveraceae

{{Short description|Family of flowering plants}} {{Redirect |Poppy family|the pop group |The Poppy Family}} {{Use dmy dates|date=February 2014}} {{Automatic taxobox | image = 2010-06-29 (28) Papaver rhoeas.jpg | image_caption = Corn poppy (''Papaver rhoeas'') | taxon = Papaveraceae | authority = Juss.{{sfn|Angiosperm Phylogeny Group|2009}} | subdivision_ranks = Genera | subdivision = See text }}

The '''Papaveraceae''' ({{IPAc-en|p|ə|ˌ|p|æ|v|ə|ˈ|r|eɪ|s|i|.|iː|,_|-|ˌ|aɪ}}),<ref>{{cite Merriam-Webster|Papaveraceae}}</ref> informally called the '''poppy family''', are an economically important family of about 42 genera and approximately 775 known species{{sfn|Christenhusz|Byng|2016}} of flowering plants in the order Ranunculales. The family is cosmopolitan, occurring in temperate and subtropical climates (mostly in the northern hemisphere) like Eastern Asia as well as California in North America. It is almost unknown in the tropics. Most are herbaceous plants, but a few are shrubs and small trees. The family currently includes two groups that have been considered to be separate families: Fumariaceae and Pteridophyllaceae. ''Papaver'' is the classical name for poppy in Latin.<ref>{{cite book |doi=10.1515/9780691209135 |title=The Gardener's Botanical |date=2020 |last1=Bayton |first1=Ross |isbn=978-0-691-20913-5 }}{{pn|date=October 2024}}</ref>

== Description == Papaveraceae are known for diverse and colorful flowers with distinctive sepals. The plants may be annual, biennial, or perennial. Usually herbaceous, a few species form shrubs or evergreen trees. All parts contain a well-developed system of latex ducts called "laticifers", that produce milky latex, a watery white, yellow or red juice.{{cn|date=June 2025}}

The leaves are alternate or sometimes whorled and have petioles and no stipules. They are usually lobed or pinnatifid (i.e. consisting of several not entirely separate leaflets), or much divided.{{cn|date=June 2025}}

The plants are hermaphroditic and pollinated mostly by insects (entomophilous), but nectaries are lacking. A few are wind pollinated (anemophilous). There is a distinct calyx and corolla, except in ''Macleaya'' where the corolla is lacking. The flowers are medium-sized or large. The terminal flowers are solitary in many species. In others the terminal inflorescence is cymose or racemose. The flowers are odourless and regular.{{cn|date=June 2025}}

There are many stamens, mostly 16 to 60, arranged in two separate whorls, the outer one with stamens alternating with petals, the inner one opposite, or numerous in the subfamily Papaveroideae.{{sfn|Stace|2010}}{{rp|86}} The gynoecium consists of a compound pistil with 2 to 100 carpels. The ovary is superior and unilocular. The ovary is either stemless (sessile) or on a short stem (stipitate). The sepals of the plant typically number half of the petals for example two sepals accompany 4 petals or 3 sepals accompany 6 petals. The pistils and stamens are hidden inside the petals.{{cn|date=June 2025}}

The non-fleshy fruit is usually a capsule, breaking open at maturity to release the seeds through pores (poricidal), through the partitions between the cells (septicidal), or by means of valves (valvular). The numerous seeds are small. Their nutritive tissue (endosperm) is oily and farinose. The fruit of ''Platystemon'' is a schizocarp.{{cn|date=June 2025}}

The basic chromosome number, ''x'', is 6, 7, 8, 9, 10 and 11, up to ''12n'' = 84 (dodecaploidy) in species of ''Papaver'', ''Argemone'' and ''Meconopsis''.{{cn|date=June 2025}}

The Papaveraceae family includes many plants that produce alkaloids, including opium poppy (''Papaver somniferum''). Opium is derived from the latex of the opium poppy seed pods and has been used for centuries due to its psychoactive properties. The main alkaloids found in opium, such as morphine, and codeine have huge impact on pharmaceuticals and a big importance on many countries that produce it on a large-scale including Afghanistan. Afghanistan has an economic dependence on opium cultivation making it hard to stop large scale production of these flowers ultimately increasing illegal production. Researchers are understanding how alkaloids are made in poppy plants to develop poppy plants with specific alkaloid levels. There is high genetic variability among poppy cultivars and environmental factors like wounding and methyl jasmonate treatment induce higher alkaloid production. Challenges still remain in understanding what effects alkaloid production for pharmaceutical purposes, highlighting importance of research in this field.{{citation needed|date=May 2024}}

== Taxonomy == The APG III system (2009; unchanged from the APG II system of 2003 and the APG system of 1998) places the family in the order Ranunculales, in the clade eudicots.{{sfn|Angiosperm Phylogeny Group|2009}} The Papaveraceae differ from the rest of the Ranunculales in some important characteristics but they share others such as the presence of isoquinoline-derived alkaloids. Based on molecular and morphological data, the family forms a clade with the families Lardizabalaceae, Circaeasteraceae, Menispermaceae, Berberidaceae and Ranunculaceae.<ref name=APW>{{Cite web |last=Stevens |first=P.F. |author-link=Peter F. Stevens |title=Ranunculales |work=Angiosperm Phylogeny Website |url=http://www.mobot.org/MOBOT/research/APweb/orders/ranunculalesweb.htm#Ranunculales |access-date=5 September 2013}}</ref>

=== Genera === [[File:HKKk9.jpg|alt=Horned poppy (Glaucium sp.)|thumb|Horned poppy (''Glaucium'' sp.)]] [[File:UK 20230418 114354.jpg|alt=Papaver sp.|thumb|''Roemeria'' sp.]] The broad circumscription of Papaveraceae in the APG III system includes three taxa that have previously been separated into different families: the Papaveraceae ''sensu stricto'', the Fumariaceae and the Pteridophyllaceae.{{sfn|Angiosperm Phylogeny Group|2009}} Thus the Cronquist system of 1981 recognised the Fumariaceae as a separate family, despite their close phylogenetic relationship to the Papaveraceae ''sensu stricto''. The three former families may be treated as subfamilies. One morphological and molecular study concluded that the former family Pteridophyllaceae has a basal position with a subsequent division into two terminal clades each containing one of the subfamilies Fumarioideae and Papaveroideae, which are clearly monophyletic.{{sfn|Hoot|Kadereit|Blattner|Jork|1997}} A more recent study includes the former Pteridophyllaceae in the Fumarioideae, dividing the Papaveraceae into only two subfamilies.{{sfn|Wang|Lu|Ren|Endress|2009}}

The internal division of the Fumarioideae shown below follows Lidén (1993),{{sfn|Lidén|1993a}} with the exception of the placement of ''Pteridophyllum''.{{sfn|Wang|Lu|Ren|Endress|2009}}<ref name=APW/> The subtribes are given by the Germplasm Resources Information Network.<ref>{{Cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/family.pl?817 |title=''Papaveraceae'' Juss., nom. cons. |work=GRIN Taxonomy for Plants |access-date=21 April 2015 |archive-url=https://web.archive.org/web/20150930063456/http://www.ars-grin.gov/cgi-bin/npgs/html/family.pl?817 |archive-date=30 September 2015 }}</ref> The division of the Papaveroideae follows Hoot et al. (1997).{{sfn|Hoot|Kadereit|Blattner|Jork|1997}} In the latter study, the tribe Eschscholzieae would be the basal clade and sister group to the rest of the subfamily, which is divided into a different terminal clade (Chelidonieae) and into its sister group, formed by the Papavereae and Platystemoneae, whose separation is not based on the data presented by these authors. For discussions of subfamilies, see Carolan et al. (2006){{sfn|Carolan|Hook|Chase|Kadereit|2006}} and Blattner & Kadereit (1999).{{sfn|Blattner|Kadereit|1999}}

====Fumarioideae==== * Subfamily Fumarioideae Eaton :*Tribe Hypecoeae Dumort. ::*''Hypecoum'' L. – Mediterranean region to Mongolia and Western China. ::*''Pteridophyllum'' Siebold & Zucc. – Japan :*Tribe {{vanchor|Fumarieae}} Dumort. ::*Subtribe {{vanchor|Corydalinae}} :::*''Adlumia'' Raf. ex DC. – Eastern North America, Korea, China :::*''Capnoides'' Mill. – Northern North America :::*''Corydalis'' DC. nom. cons. – Eurasia, North America, East Africa :::*''Dactylicapnos'' Wall. – Himalayas :::*''Dicentra'' Bernh. nom. cons. – Eastern Asia, North America :::*''Ehrendorferia'' Fukuhara & Lidén – Western United States :::*''Ichtyoselmis'' Lidén & Fukuhara – China :::*''Lamprocapnos'' Endl. – China, Korea ::*Subtribe {{vanchor|Fumariinae}} :::*''Ceratocapnos'' Durieu – South west of Europe, north west of Africa :::*''Cryptocapnos'' Rech.f. – Central Afghanistan :::*''Cysticapnos'' Mill. – South Africa :::*''Discocapnos'' Cham. & Schltdl. – South Africa :::*''Fumaria'' L. – Mediterranean region, Himalayas, East Africa :::*''Fumariola'' Korsh. – Central Asia :::*''Platycapnos'' (DC.) Bernh. – Western Mediterranean region :::*''Pseudofumaria'' Medik. – Italy, Balkans :::*''Rupicapnos'' Pomel – North west Africa :::*''Sarcocapnos'' DC. – Spain, Morocco, Algeria :::*''Trigonocapnos'' Schltr. – South Africa

<gallery caption="Photos of Fumarioideae (the fumewort subfamily)"> Hypecoum procumbens FlowerCloseup 25April2009 CampodeCalatrava.jpg|''Hypecoum procumbens'' Corydalisaurea.jpg|Scrambled eggs (''Corydalis aurea'') Corydalis solida01.jpg|Fumewort (''Corydalis solida'') Corydalis-cheilanthifolia-habitus.jpg|Ferny corydalis (''Corydalis cheilanthifolia'') Pale corydalis closeup.jpg|Pale corydalis (''Capnoides sempervirens'' or ''Corydalis sempervirens'') 3949 - Dicentra scandens (Kletterndes Tränendes Herz).JPG|Climbing bleeding-heart (''Dactylicapnos scandens'' or ''Dicentra scandens'') Dicentra cucullaria.png|Dutchman's breeches (''Dicentra cucullaria'') Fringed bleeding-heart flower cluster.jpg|Fringed bleeding-heart (''Dicentra eximia'') Dicentra peregrina (flower).JPG|Komakusa (''Dicentra peregrina'') Dicentrachrysantha.jpg|Golden eardrops (''Ehrendorferia chrysantha'' or ''Dicentra chrysantha'') Tränendes Herz (Dicentra spectabilis).jpg|Asian bleeding-heart (''Lamprocapnos spectabilis'' or ''Dicentra spectabilis'') Earth smoke 04.jpg|Earth smoke (''Fumaria officinalis'') Corydalis lutea Flowers.JPG|Yellow corydalis (''Pseudofumaria lutea'' or ''Corydalis lutea'') Sarcocapnos enneaphylla733.jpg|''Zapatitos de la Virgen'' (''Sarcocapnos enneaphylla'') </gallery>

====Papaveroideae==== * Subfamily Papaveroideae Eaton :*Tribe {{vanchor|Eschscholzieae}} Baill. ::*''Dendromecon'' Benth. – California. ::*''Eschscholzia'' Cham. – Western North America. ::*''Hunnemannia'' Sweet – Eastern Mexico. :*Tribe {{vanchor|Chelidonieae}} Dumort. ::*''Bocconia'' L. – Central and southern America, Antilles ::*''Chelidonium'' L. – Eurasia ::*''Coreanomecon'' Nakai – Korea ::*''Dicranostigma'' Hook.f. & Thomson – Central Asia ::*''Eomecon'' Hance – Eastern China ::*''Glaucium'' Mill. – Europe to Central Asia ::*''Hylomecon'' Maxim. – Eastern Asia ::*''Macleaya'' R.Br. – Eastern Asia ::*''Sanguinaria'' L. – Eastern North America ::*''Stylophorum'' Nutt. – Eastern North America, Eastern Asia :*Tribe {{vanchor|Platystemoneae}} Spach ::*''Hesperomecon'' Greene – Western North America ::*''Meconella'' Nutt. – Western North America ::*''Platystemon'' Benth. – Western North America :*Tribe {{vanchor|Papavereae}} Dumort. ::*''Arctomecon'' Torr. & Frém. – Western North America ::*''Argemone'' L. – North America, Antilles, central and southern America, Hawaii ::*''Canbya'' Parry – Western North America ::*''Cathcartia'' Hook.f. – China and the Himalayas, split from ''Meconopsis'' ::*''Meconopsis'' Vig. – Central southern Asia, western Europe; paraphyletic{{sfn|Kadereit|Schwarzbach|Jork|1997}} ::*''Papaver'' L. – Northern hemisphere, South Africa, Cape Verde; paraphyletic{{sfn|Kadereit|Schwarzbach|Jork|1997}} ::*''Roemeria'' Medik. – Mediterranean region, south west Asia ::*''Romneya'' Harv. – California ::*''Stylomecon'' G. Taylor – California

<gallery caption="Photos of Papaveroideae (the poppy subfamily)"> Flickr - brewbooks - Island Tree Poppy (1).jpg|Channel Island tree poppy (''Dendromecon harfordii'') Hunnemannia fumariifolia flowers 2002-10-10.jpg|Tulip poppy (''Hunnemannia fumariifolia'') Starr 070308-5399 Bocconia frutescens.jpg|Plume poppy (''Bocconia frutescens'') Flower October 2008-1.jpg|Greater celandine (''Chelidonium majus'') Dicranostigma erectum2.JPG|Eastern horned poppy (''Dicranostigma erectum'') Ab plant 60.jpg|Red horned poppy (''Glaucium corniculatum'') Eomecon chionantha.jpg|Snow-poppy (''Eomecon chionantha'') Hylomecon japonica 01.JPG|Forest poppy (''Hylomecon vernalis'') Bloodroot (Sanguinaria canadensis) - Guelph.jpg|Bloodroot (''Sanguinaria canadensis'') Stylophorum diphyllum flowers cropped.png|Wood poppy (''Stylophorum diphyllum'') Platystemon californicus (1).jpg|Creamcups (''Platystemon californicus'') Arctomecon californica 3.jpg|Bearpaw poppy (''Arctomecon californica'') White poppy Argemone munita close.jpg|Flat-bud prickly poppy (''Arctomecon munita'') Pygmy Poppy, Canbya candida.jpg|Pygmy poppy (''Canbya candida'') Arctic poppies at Showa Memorial Park.jpg|Arctic poppy (''Papaver nudicaule'') Papaver(Kirechko).jpg|Persian poppy (''Papaver bracteatum'') Papaver somniferum (Kowal).jpg|Opium poppy (''Papaver somniferum'') Romneya-coulteri-habitus.jpg|Matilija poppy (''Romneya coulteri'') </gallery>

== Ecology == [[File:Pteridophyllum racemosum.JPG|thumb|upright=0.8|Fern-poppy (''Pteridophyllum racemosum'')]] Pollination is entomophile (basically by flies and wasps and bees, less often by beetles), except in ''Bocconia'' and ''Macleaya''. In Papaveroideae, the reward is pollen as there is no nectar. The visual attractant is the petals that are usually brightly coloured and often have basal guides, sometimes the attractant can also be the androecium as the petals do not last long. Some species, mostly those from the arctic and alpine regions, reinforce their attraction with floral fragrance (for example, ''Papaver alpinum'' smells of cloves), which in the case of ''Romneya'' drugs the insects. The anthers and stigmas mature at the same time, but ''Bocconia'' is clearly protogynous, the stigmas emerge from the calyx that encloses them. Autopollination is common and in some cases (for example, ''Roemeria hybrida'') it occurs before the bud opens (cleistogamy). The presence of an aril suggests dispersion of seeds by ants (myrmecochory), once they have been expelled by the fruit. In the case of ''Bocconia'' the seeds remain attached to the replums after the capsule's valves have fallen leaving their brilliant red or orange arils exposed, which attract birds to feed on them, facilitating their dispersal (ornithochory). Seeds that lack an aril appear to be dispersed by the wind (anemochory) for capsules that open, in the other cases they are freed when the fruit decomposes. Many Fumarioideae species have explosive fruits (ballistic), while ''Rupicapnos'' and ''Sarcocapnos'' species are chasmophytes, growing on rocks, and their fruit's peduncles and pedicels are geotropic and they lengthen so that the seeds bury into the base of the plant.{{cn|date=June 2025}}

The Papaveroideae typically grow in cooler and wooded areas, forming part of the undergrowth. They have adapted to arctic and alpine habitats and to arid, Mediterranean areas, many species are ruderal and segetal (growing in cornfields). ''Pteridophyllum'' grows in the undergrowth of woods of needle-leaved trees between {{convert|1,000|and|2,000|m}}. The Fumarioideae are basically found in open, rocky, alpine landscapes or vertical or overhanging cracks, while some species are ruderal or segetal.{{cn|date=June 2025}}

== Phytochemistry == [[File:Argemone maxicana in Hyderabad, AP W IMG 1102.jpg|thumb|upright=0.8|Mexican prickly poppy (''Argemone mexicana'')]] Alkaloids: The isoquinolinic alkaloids present in the family are well known. They are derived from berberine, tetrahydroberberine, protopine and benzophenanthridine in Papaveroideae, and from spirobenzylisoquinoline and cularine in Fumarioideae, as well as from other groups that give them pharmacological properties: derivatives of aporphine, morphinan, pavine, isopavine, narceine and rhoeadine.{{cn|date=June 2025}}

Others: Other characteristic substances contained within these species include: meconic acid and chelidonic acid, as well as cyanogenic glycoside compounds derived from tyrosine: dhurrin and triglochinin; in the Fumarioideae while the Chelidonieae contain the free amino acid δ-acetylornithine.{{cn|date=June 2025}}

Flavonoids: Iridoids and proanthocyanidins absent. Flavonols, kaempferol and/or quercetin present.{{cn|date=June 2025}}

Many of these plants are poisonous. The Mexican prickly poppy is poisonous if taken internally and may cause oedema and glaucoma. Even if an animal, such as a goat, should persist in grazing on this plant, not only will the animal suffer but so will those who drink its milk, because the poisons are passed along in the milk.{{cn|date=June 2025}}

==Fossil record== thumb|''Paleoaster'' sp. The fossils of the late Cretaceous poppy ''Palaeoaster inquirenda'' from the Western Interior of North America occurs from 74.5-million-year-old deposits in the Fruitland Formation in New Mexico to 64.5-million-year-old deposits in the Hell Creek Formation in North Dakota. Dehiscent fruit fossils of ''Palaeoaster'' have been found at the excavation site for the well-known ''Tyrannosaurus rex'' specimen BHI 3033. The seed capsule of ''Palaeoaster'' has some similarities to that of the extant poppy genus ''Romneya''.{{sfn|Smith|2001}}

''Papaverites'', a fossil fruit from the Eocene of Germany, may be associated with Papaveraceae.{{sfn|Friedel|1927}} Chesters et al. (1967) mentions ''Papaver pictum'' from the Oligocene of England.{{sfn|Willemstein|1987}}

== Cultivation == {{multiple image |orientiation=horizontal |total_width=330 |image1=California Poppy Eschscholzia californica 02.jpg |caption1=California poppy (''Eschscholzia californica'') |image2=Himalayan Blue Poppy - Meconopsis betonicifolia - ヒマラヤの青いケシ (8932550724) (2).jpg |caption2=Himalayan blue poppy (''Meconopsis betonicifolia'') }} The family is well known for its striking flowers, with many species grown as ornamental plants, including California poppy (''Eschscholtzia californica'', the California state flower), the stunning blue Himalayan poppies (''Meconopsis''), several species of ''Papaver'', and the wildflower bloodroot. Only two species are of economic importance for the production of opium and its derivatives for pharmaceutical use: ''Papaver somniferum'' is cultivated legally in order to obtain morphine and other opiates, and ''Papaver bracteatum'', for thebaine. ''Papaver somniferum'' is also the source of the poppy seeds used in cooking and baking, and poppy seed oil. The illegal cultivation of poppies in Asia for the production of opium and heroin is virtually equal to the legal production in the rest of the world. Some Funarioideae have a limited use in gardening, with ''Lamprocapnos spectabilis'' ("bleeding heart"), and ''Pseudofumaria lutea'' ("yellow corydalis") commonly used. Chinese traditional medicine used the boiled and dried tubers of ''Corydalis yanhusuo'' ("yanhusuo").{{cn|date=June 2025}}

==Symbolism== {{Main|Poppy}}

The opium poppy and corn poppy are symbols, respectively, of sleep and death. In Great Britain, Canada, the United States, and Australia the corn poppy is worn in remembrance of World War I.

== References == {{Reflist|24em}}

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== External links ==

{{Commons category}} * [http://www.topwalks.net/plants/generos/papaveraceae_01.htm Papaveraceae in Topwalks] * [http://delta-intkey.com/angio/www/papavera.htm Papaveraceae] {{Webarchive|url=https://web.archive.org/web/20060320121514/http://delta-intkey.com/angio/www/papavera.htm |date=20 March 2006 }} [sensu stricto] in L. Watson and M.J. Dallwitz (1992 onwards). [http://delta-intkey.com/angio/ ''The families of flowering plants''] {{Webarchive|url=https://web.archive.org/web/20070103200438/http://delta-intkey.com/ |date=3 January 2007 }}. * Papaveraceae [sensu lato] in the [http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=10650 ''Flora of North America''] * [http://www.csdl.tamu.edu/FLORA/cgi/gateway_family?fam=Papaveraceae links at CSDL] {{Webarchive|url=https://web.archive.org/web/20080203152151/http://www.csdl.tamu.edu/FLORA/cgi/gateway_family?fam=Papaveraceae |date=3 February 2008 }} * [http://flowersinisrael.com/FamPapaveraceae.html Family Papaveraceae] {{Webarchive|url=https://web.archive.org/web/20121011053907/http://flowersinisrael.com/FamPapaveraceae.html |date=11 October 2012 }} Flowers in Israel * [http://www.mobot.org/MOBOT/Research/APweb/maps/pap-pterido.gif Mapa de Pteridophylloideae] * [http://www.mobot.org/MOBOT/Research/APweb/maps/pap-pap.gif Mapa de Papaveroideae] * [http://www.mobot.org/MOBOT/Research/APweb/maps/pap-fum.gif Mapa de Fumarioideae]

Category:Papaveraceae Category:Eudicot families Category:Taxa named by Antoine Laurent de Jussieu