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The Papaveraceae, informally known as the poppy family, are an economically important family of about 42 and approximately 775 known of in the order . The family is cosmopolitan, occurring in and climates (mostly in the northern hemisphere) like Eastern Asia as well as California in North America. It is almost unknown in the . Most are , but a few are and small . 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.

(2025). 9780691209135


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

The are alternate or sometimes whorled and have petioles and no . They are usually lobed or pinnatifid (i.e. consisting of several not entirely separate leaflets), or much divided.

The plants are and mostly by (), but nectaries are lacking. A few are wind pollinated (). There is a distinct calyx and corolla, except in where the corolla is lacking. The flowers are medium-sized or large. The terminal flowers are solitary in many species. In others the terminal is cymose or . The flowers are odourless and regular.

There are many , 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 . The gynoecium consists of a compound with 2 to 100 carpels. The ovary is superior and . 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.

The non-fleshy is usually a capsule, breaking open at maturity to release the 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 is a .

The basic , x, is 6, 7, 8, 9, 10 and 11, up to 12n = 84 (dodecaploidy) in species of , and .

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.


Taxonomy
The APG III system (2009; unchanged from the APG II system of 2003 and the of 1998) places the family in the order , in the clade . 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 , , , and .


Genera
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. Thus the 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. A more recent study includes the former Pteridophyllaceae in the Fumarioideae, dividing the Papaveraceae into only two subfamilies.

The internal division of the Fumarioideae shown below follows Lidén (1993), with the exception of the placement of Pteridophyllum. The subtribes are given by the Germplasm Resources Information Network. The division of the Papaveroideae follows Hoot et al. (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) and Blattner & Kadereit (1999).


Fumarioideae
*Tribe Hypecoeae Dumort.
:* – Mediterranean region to Mongolia and Western China.
:* Siebold & Zucc. – Japan
*Tribe Dumort.
:*Subtribe
::* Raf. ex DC. – Eastern North America, Korea, China
::* – Northern North America
::* DC. nom. cons. – Eurasia, North America, East Africa
::* Wall. – Himalayas
::* Bernh. nom. cons. – Eastern Asia, North America
::* Fukuhara & Lidén – Western United States
::* Lidén & Fukuhara – China
::* Endl. – China, Korea
:*Subtribe
::* Durieu – South west of Europe, north west of Africa
::* Rech.f. – Central Afghanistan
::* Mill. – South Africa
::* Cham. & Schltdl. – South Africa
::* L. – Mediterranean region, Himalayas, East Africa
::* Korsh. – Central Asia
::* (DC.) Bernh. – Western Mediterranean region
::* Medik. – Italy, Balkans
::* – North west Africa
::* DC. – Spain, Morocco, Algeria
::* Schltr. – South Africa

Hypecoum procumbens FlowerCloseup 25April2009 CampodeCalatrava.jpg| Hypecoum procumbens Corydalisaurea.jpg| ( Corydalis aurea) Corydalis solida01.jpg|Fumewort ( Corydalis solida) Corydalis-cheilanthifolia-habitus.jpg|Ferny corydalis ( Corydalis cheilanthifolia) Pale corydalis closeup.jpg| ( Capnoides sempervirens or Corydalis sempervirens) 3949 - Dicentra scandens (Kletterndes Tränendes Herz).JPG| ( Dactylicapnos scandens or Dicentra scandens) Dicentra cucullaria.png|Dutchman's breeches ( Dicentra cucullaria) Fringed bleeding-heart flower cluster.jpg| ( Dicentra eximia) Dicentra peregrina (flower).JPG|Komakusa ( Dicentra peregrina) Dicentrachrysantha.jpg|Golden eardrops ( Ehrendorferia chrysantha or Dicentra chrysantha) Tränendes Herz (Dicentra spectabilis).jpg| ( 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| ( Sarcocapnos enneaphylla)


Papaveroideae
*Tribe Baill.
:* – California.
:* Cham. – Western North America.
:* Sweet – Eastern Mexico.
*Tribe Dumort.
:* Bocconia L. – Central and southern America, Antilles
:* L. – Eurasia
:* Nakai – Korea
:* Hook.f. & Thomson – Central Asia
:* Hance – Eastern China
:* Mill. – Europe to Central Asia
:* – Eastern Asia
:* R.Br. – Eastern Asia
:* L. – Eastern North America
:* – Eastern North America, Eastern Asia
*Tribe Spach
:* Greene – Western North America
:* Nutt. – Western North America
:* – Western North America
*Tribe Dumort.
:* & Frém. – Western North America
:* L. – North America, Antilles, central and southern America, Hawaii
:* Parry – Western North America
:* Hook.f. – China and the Himalayas, split from Meconopsis
:* Vig. – Central southern Asia, western Europe; paraphyletic
:* L. – Northern hemisphere, South Africa, Cape Verde; paraphyletic
:* Medik. – Mediterranean region, south west Asia
:* Harv. – California
:* G. Taylor – California

Flickr - brewbooks - Island Tree Poppy (1).jpg|Channel Island tree poppy ( Dendromecon harfordii) Hunnemannia fumariifolia flowers 2002-10-10.jpg| ( Hunnemannia fumariifolia) Starr 070308-5399 Bocconia frutescens.jpg|Plume poppy ( Bocconia frutescens) Flower October 2008-1.jpg| ( Chelidonium majus) Dicranostigma erectum2.JPG| ( Dicranostigma erectum) Ab plant 60.jpg| ( Glaucium corniculatum) Eomecon chionantha.jpg| ( Eomecon chionantha) Hylomecon japonica 01.JPG| ( Hylomecon vernalis) Bloodroot (Sanguinaria canadensis) - Guelph.jpg| ( Sanguinaria canadensis) Stylophorum diphyllum flowers cropped.png|Wood poppy ( Stylophorum diphyllum) Platystemon californicus (1).jpg| ( Platystemon californicus) Arctomecon californica 3.jpg|Bearpaw poppy ( Arctomecon californica) White poppy Argemone munita close.jpg| ( Arctomecon munita) Pygmy Poppy, Canbya candida.jpg| ( 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| ( Romneya coulteri)


Ecology
Pollination is (basically by and , less often by ), except in Bocconia and . 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 , sometimes the attractant can also be the 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, smells of cloves), which in the case of drugs the insects. The and stigmas mature at the same time, but Bocconia is clearly , the stigmas emerge from the calyx that encloses them. is common and in some cases (for example, Roemeria hybrida) it occurs before the bud opens (). The presence of an suggests dispersion of seeds by ants (), once they have been expelled by the fruit. In the case of Bocconia the seeds remain attached to the 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 (). Seeds that lack an aril appear to be dispersed by the wind () for capsules that open, in the other cases they are freed when the fruit decomposes. Many species have explosive fruits (ballistic), while and species are , growing on rocks, and their fruit's peduncles and pedicels are and they lengthen so that the seeds bury into the base of the plant.

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 and segetal (growing in cornfields). grows in the undergrowth of woods of needle-leaved trees between . The Fumarioideae are basically found in open, rocky, alpine landscapes or vertical or overhanging cracks, while some species are ruderal or segetal.


Phytochemistry
: The alkaloids present in the family are well known. They are derived from , tetrahydroberberine, and benzophenanthridine in Papaveroideae, and from spirobenzylisoquinoline and cularine in Fumarioideae, as well as from other groups that give them pharmacological properties: derivatives of , , pavine, , and .

Others: Other characteristic substances contained within these species include: and , as well as cyanogenic glycoside compounds derived from : and triglochinin; in the Fumarioideae while the Chelidonieae contain the free amino acid δ-acetylornithine.

: and absent. , and/or present.

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


Fossil record
The fossils of the late poppy Palaeoaster inquirenda from the Western Interior of North America occurs from 74.5 million year old deposits in the Fruitland Formation in to 64.5 million year old deposits in the Hell Creek Formation in . 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 .

Papaverites, a fossil fruit from the of , may be associated with Papaveraceae. Chesters et al. (1967) mentions Papaver pictum from the of .


Cultivation
The family is well known for its striking flowers, with many species grown as , including ( Eschscholtzia californica, the California ), the stunning blue ( Meconopsis), several species of Papaver, and the wildflower . Only two species are of economic importance for the production of and its derivatives for pharmaceutical use: Papaver somniferum is cultivated legally in order to obtain and other , and Papaver bracteatum, for . Papaver somniferum is also the source of the used in and , and poppy seed oil. The illegal cultivation of poppies in Asia for the production of opium and 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").


Symbolism
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.


Bibliography


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