Papaveraceae

Papaveraceae
	Corn poppy ( Papaver rhoeas )
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Order:	Ranunculales
Family:	Papaveraceae ; Juss.
Genera
	See text

Last updated April 27, 2024

The Papaveraceae /pəˌpævəˈreɪsiˌiː/ ^[2] are an economically important family of about 42 genera and approximately 775 known species ^[3] of flowering plants in the order Ranunculales, informally known as the poppy family. 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.^[4]

Description

This family is known for its diverse and colorful flowers and distinctive sepals. The plants may be annual, biennial, or perennial. Usually herbaceous, a few spices form shrubs or evergreen trees. They are laticiferous, producing latex, which may be milky or watery, coloured or plain. All parts contain a well-developed duct system (these ducts are called "laticifers"), producing a milky latex, a watery white, yellow or red juice.

The simple leaves are alternate or sometimes whorled. They have petioles and are not enclosed by a sheath. The leaves are usually lobed or pinnatifid (i.e. consisting of several not entirely separate leaflets), or much divided. There are no stipules.

The plants are hermaphroditic and are pollinated mostly by insects (entomophilous); flower 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.

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.^[5]^: 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 Stamen are hidden inside the petals.

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.

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 .

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. Papaver somniferum is also found to have anticancer properties mainly against cervical and colon cancers as it was found to disrupt cell membrane of tumor cells. The opium poppy (Papaver somniferum) shares a common ancestor with Macleya cordata, and Eschschloizia californica. Papaver somniferum evolved from Aquilegia coerulea then Nelumbo nucifera and Macadamia intergrifolia. The furthest known ancestor to Papaver somniferum is Oryza sativa.

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.^[1] 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.^[6]

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.^[1] 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.^[7] A more recent study includes the former Pteridophyllaceae in the Fumarioideae, dividing the Papaveraceae into only two subfamilies.^[8]

The internal division of the Fumarioideae shown below follows Lidén (1993),^[9] with the exception of the placement of Pteridophyllum.^[8]^[6] The subtribes are given by the Germplasm Resources Information Network.^[10] The division of the Papaveroideae follows Hoot et al. (1997).^[7] 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)^[11] and Blattner & Kadereit (1999).^[12]

Fumarioideae

Subfamily Fumarioideae Eaton

Tribe Hypecoeae Dumort.

Hypecoum L. – Mediterranean region to Mongolia and Western China.
Pteridophyllum Siebold & Zucc. – Japan

Tribe FumarieaeDumort.

Subtribe 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 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

Photos of Fumarioideae (the fumewort subfamily)
Hypecoum procumbens
Scrambled eggs (Corydalis aurea)
Fumewort (Corydalis solida)
Ferny corydalis (Corydalis cheilanthifolia)
Pale corydalis (Capnoides sempervirens)
Climbing bleeding-heart (Dactylicapnos scandens or Dicentra scandens)
Dutchman's breeches (Dicentra cucullaria)
Fringed bleeding-heart (Dicentra eximia)
Komakusa (Dicentra peregrina)
Golden eardrops (Ehrendorferia chrysantha or Dicentra chrysantha)
Asian bleeding-heart (Lamprocapnos spectabilis or Dicentra spectabilis)
Earth smoke (Fumaria officinalis)
Yellow corydalis (Pseudofumaria lutea or Corydalis lutea)
Zapatitos de la Virgen (Sarcocapnos enneaphylla)

Papaveroideae

Subfamily Papaveroideae Eaton

Tribe Eschscholzieae Baill.

Dendromecon Benth. – California.
Eschscholzia Cham. – Western North America.
Hunnemannia Sweet – Eastern Mexico.

Tribe ChelidonieaeDumort.

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 Platystemoneae Spach

Hesperomecon Greene – Western North America
Meconella Nutt. – Western North America
Platystemon Benth. – Western North America

Tribe PapavereaeDumort.

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^[13]
Papaver L. – Northern hemisphere, South Africa, Cape Verde; paraphyletic^[13]
Roemeria Medik. – Mediterranean region, south west Asia
Romneya Harv. – California
Stylomecon G. Taylor – California

Photos of Papaveroideae (the poppy subfamily)
Channel Island tree poppy (Dendromecon harfordii)
Tulip poppy (Hunnemannia fumariifolia)
Plume poppy (Bocconia frutescens)
Greater celandine (Chelidonium majus)
Eastern horned poppy (Dicranostigma erectum)
Red horned poppy (Glaucium corniculatum)
Snow-poppy (Eomecon chionantha)
Forest poppy (Hylomecon vernalis)
Bloodroot (Sanguinaria canadensis)
Wood poppy (Stylophorum diphyllum)
Creamcups (Platystemon californicus)
Bearpaw poppy (Arctomecon californica)
Flat-bud prickly poppy (Arctomecon munita)
Pygmy poppy (Canbya candida)
Arctic poppy (Papaver nudicaule)
Persian poppy (Papaver bracteatum)
Opium poppy (Papaver somniferum)
Matilija poppy (Romneya coulteri)

Ecology

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.

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 1,000 and 2,000 metres (3,300 and 6,600 ft). The Fumarioideae are basically found in open, rocky, alpine landscapes or vertical or overhanging cracks, while some species are ruderal or segetal.

Phytochemistry

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.

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.

Flavonoids: Iridoids and proanthocyanidins absent. Flavonols, kaempferol and/or quercetin present.

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.

Fossil record

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 .^[14]

Papaverites, a fossil fruit from the Eocene of Germany, may be associated with Papaveraceae.^[15] Chesters et al. (1967) mentions Papaver pictum from the Oligocene of England.^[16]

Cultivation

California poppy (Eschscholzia californica)

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").

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.

Related Research Articles

Eschscholzia californica, the California poppy, golden poppy, California sunlight or cup of gold, is a species of flowering plant in the family Papaveraceae, native to the United States and Mexico. It is cultivated as an ornamental plant flowering in summer, with showy cup-shaped flowers in brilliant shades of red, orange and yellow. It is also used as food or a garnish. It became the official state flower of California in 1903.

A poppy is a flowering plant in the subfamily Papaveroideae of the family Papaveraceae. Poppies are herbaceous plants, often grown for their colourful flowers. One species of poppy, Papaver somniferum, is the source of the narcotic drug mixture opium which contains powerful medicinal alkaloids such as morphine and has been used since ancient times as an analgesic and narcotic medicinal and recreational drug. It also produces edible seeds. Following the trench warfare in the poppy fields of Flanders, Belgium during World War I, poppies have become a symbol of remembrance of soldiers who have died during wartime, especially in the UK, Canada, Australia, New Zealand and other Commonwealth realms.

Ranunculales is an order of flowering plants. Of necessity it contains the family Ranunculaceae, the buttercup family, because the name of the order is based on the name of a genus in that family. Ranunculales belongs to a paraphyletic group known as the basal eudicots. It is the most basal clade in this group; in other words, it is sister to the remaining eudicots. Widely known members include poppies, barberries, hellebores, and buttercups.

Fumarioideae is a subfamily of the family Papaveraceae. It was formerly treated as a separate family, the Fumariaceae. It consists of about 575 species of herbaceous plants in 20 genera, native to the Northern Hemisphere and South Africa. The largest genus is Corydalis.

Papaver rhoeas, with common names including common poppy, corn poppy, corn rose, field poppy, Flanders poppy, and red poppy, is an annual herbaceous species of flowering plant in the poppy family Papaveraceae. It is native to north Africa and temperate Eurasia and is introduced into temperate areas on all other continents except Antarctica.

Papaver is a genus of 70–100 species of frost-tolerant annuals, biennials, and perennials native to temperate and cold regions of Eurasia, Africa and North America. It is the type genus of the poppy family, Papaveraceae.

Papaver somniferum, commonly known as the opium poppy or breadseed poppy, is a species of flowering plant in the family Papaveraceae. It is the species of plant from which both opium and poppy seeds are derived and is also a valuable ornamental plant grown in gardens. Its native range was east of the Mediterranean Sea, but now is obscured by ancient introductions and cultivation, being naturalized across much of Europe and Asia.

Meconopsis is a genus of flowering plants in the poppy family Papaveraceae. It was created by French botanist Viguier in 1814 for the species known by the common name Welsh poppy, which Carl Linnaeus had described as Papaver cambricum. The genus name means "poppy-like". Himalayan species discovered later were also placed in Meconopsis. In the 21st century, it was discovered that the Himalayan species were less closely related to the Welsh poppy, which has been restored to Papaver. All species now placed in Meconopsis are native to the Himalayas and surrounding regions. They have attractive, usually blue flowers.

Papaver cambricum, synonym Meconopsis cambrica, the Welsh poppy, is a perennial flowering plant in the poppy family Papaveraceae. It has yellow to orange flowers and is widely grown as a garden plant. It is a native of damp, rocky sites in upland areas of Western Europe from the British Isles to the Iberian Peninsula. It has been used since 2006 as the basis for the logo of the political party Plaid Cymru.

<span class="mw-page-title-main">Poppy tea</span> Herbal tea made out of poppy straw or poppy seeds

Poppy tea is an herbal tea infusion brewed from poppy straw or seeds of several species of poppy. The species most commonly used for this purpose is Papaver somniferum, which produces opium as a natural defense against predators. In the live flower, opium is released when the surface of the bulb, called the seed pod, is pierced or scraped. For the purpose of the tea, dried pods are more commonly used than the pods of the live flower. The walls of the dried pods contain opiate alkaloids, primarily consisting of morphine.

Roemeria argemone is a species of flowering plant in the poppy family Papaveraceae. Its common names include long pricklyhead poppy, prickly poppy and pale poppy. Its native range includes parts of Eurasia and North Africa, but it can be found growing wild in parts of North America, where it is an introduced species. It is cultivated as an ornamental plant.

Meconic acid, also known as acidum meconicum and poppy acid, is a chemical substance found in certain plants of the poppy family, Papaveraceae, such as Papaver somniferum and Papaver bracteatum. Meconic acid constitutes about 5% of opium and can be used as an analytical marker for the presence of opium. Meconic acid has erroneously been described as a mild narcotic, but it has little or no physiological activity, and is not used medicinally. Meconic acid forms salts with alkaloids and metals. These salts as well as meconic acid esters are called meconates. Meconic acid was first isolated by Friedrich Sertürner in 1805.

Corydalis nobilis, the Siberian corydalis, is a perennial plant native to Siberia, Xinjiang and Kazakhstan. It was introduced to Europe by Linnaeus, who had asked his friend Erich Laxmann for seeds of Lamprocapnos spectabilis, but was sent seeds of C. nobilis instead. Both Lamprocampnos and Corydalis are members of the family Papaveraceae, with seeds having an attached elaiosome that makes them attractive to ants, which disperse the seeds.

Papaver setigerum, common name poppy of Troy or dwarf breadseed poppy, is a herbaceous annual plant of the family Papaveraceae.

<span class="mw-page-title-main">Chelidonine</span> Chemical compound

Chelidonine is an isolate of Papaveraceae with acetylcholinesterase and butyrylcholinesterase inhibitory activity.

Papaveroideae is a subfamily of the family Papaveraceae.

<span class="mw-page-title-main">Isoquinoline alkaloids</span>

Isoquinoline alkaloids are natural products of the group of alkaloids, which are chemically derived from isoquinoline. They form the largest group among the alkaloids.

<i>Papaver heterophyllum</i> Plant species

Papaver heterophyllum, previously known as Stylomecon heterophylla, and better known as the wind poppy, is a winter annual herbaceous plant. It is endemic to the western California Floristic Province and known to grow in the area starting from the San Francisco Bay Area of Central Western California southwards to northwestern Baja California, Mexico. Its main habitat is often described as mesic and shady, with loamy soils such as soft sandy loam, clay loam, and leaf mold loam.

Oreomecon is a genus in the poppy family Papaveraceae. It was established in 2022 for what was previously treated as Papaver sect. Meconella in order to ensure that the genus Papaver was monophyletic. As of June 2023, names in the genus Oreomecon had only been published for better known and phylogenetically understood species present in Europe, either as natives or aliens.

References

1 2 3 Angiosperm Phylogeny Group 2009.
↑ "Papaveraceae". Merriam-Webster.com Dictionary .
↑ Christenhusz & Byng 2016.
↑ Bayton, Ross (10 March 2020). The Gardener's Botanical. Princeton University Press. ISBN 978-0-691-20913-5.
↑ Stace 2010.
1 2 Stevens, P.F. "Ranunculales". Angiosperm Phylogeny Website. Retrieved 5 September 2013.
1 2 Hoot et al. 1997.
1 2 Wang et al. 2009.
↑ Lidén 1993a.
↑ "Papaveraceae Juss., nom. cons". GRIN Taxonomy for Plants. Archived from the original on 30 September 2015. Retrieved 21 April 2015.
↑ Carolan et al. 2006.
↑ Blattner & Kadereit 1999.
1 2 Kadereit, Schwarzbach & Jork 1997.
↑ Smith 2001.
↑ Friedel 1927.
↑ Willemstein 1987.

Bibliography

Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi: 10.1111/j.1095-8339.2009.00996.x . hdl: 10654/18083 .
Blattner, Frank R. & Kadereit, Joachim W. (1999). "Morphological evolution and ecological diversification of the forest-dwelling poppies (Papaveraceae: Chelidonioideae) as deduced from a molecular phylogeny of the ITS region". Plant Systematics and Evolution. 219 (3–4): 181–197. doi:10.1007/bf00985578. S2CID 28548382.
Carolan, James C.; Hook, Ingrid L.I.; Chase, Mark W.; Kadereit, Joachim W. & Hodkinson, Trevor R. (2006). "Phylogenetics of Papaver and Related Genera Based on DNA Sequences from ITS Nuclear Ribosomal DNA and Plastid trnL Intron and trnL–F Intergenic Spacers". Annals of Botany. 98 (1): 141–155. doi:10.1093/aob/mcl079. PMC 2803553 . PMID 16675606.
Christenhusz, M. J. M. & Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3): 201–217. doi: 10.11646/phytotaxa.261.3.1 .
Friedel, Jean (1927). "Filiation des Papavéracées". Bulletin de la Société Botanique de France (in French). 74 (4): 673–688. doi:10.1080/00378941.1927.10833678.
Hoot, S.B.; Kadereit, J.W.; Blattner, F.R.; Jork, K.B.; Schwarzbach, A.E. & Crane, P.R. (1997). "Data congruence and phylogeny of the Papaveraceae s.l. based on four data sets: atpB and rbcL sequences, trnK restriction sites, and morphological characters". Systematic Botany. 22 (3): 575–590. doi:10.2307/2419829. JSTOR 2419829.
Kadereit, J.W. (1993). "Papaveraceae". In Kubitzki, Rohwer & Bittrich (1993)
Kadereit, J.W.; Schwarzbach, A.E.; Jork, K.B. (1997). "The phylogeny of Papaver s.l. (Papaveraceae): polyphyly or monophyly?". Plant Systematics and Evolution. 204 (1–2): 75–98. doi:10.1007/bf00982533. S2CID 20019969.
Kubitzki, K.; Rohwer, J.G.; Bittrich, V., eds. (1993). The Families and Genera of Vascular Plants. II. Flowering Plants - Dicotyledons. Berlin: Springer-Verlag. ISBN 978-3-540-55509-4.
Lidén, M. (1993a). "Fumariaceae". In Kubitzki, Rohwer & Bittrich (1993)
Lidén, M. (1993b). "Pteridophyllaceae". In Kubitzki, Rohwer & Bittrich (1993)
Lidén, M.; Fukuhara, T.; Rylander, J. & Oxelman, B. (1997). "Phylogeny and classification of Fumariaceae, with emphasis on Dicentra s.l., based on the plastid gene rps16 intron". Plant Systematics and Evolution. 206 (1–4): 411–420. doi:10.1007/bf00987960. S2CID 24285424.
Smith, Una R. (2001). "Revision of the Cretaceous Fossil Genus Palaeoaster (Papaveraceae) and Clarification of Pertinent Species of Eriocaulon, Palaeoaster and Sterculiocarpus". Novon. 11 (2): 258–260. doi:10.2307/3393064. JSTOR 3393064.
Stace, C. A. (2010). New Flora of the British Isles (3rd ed.). Cambridge, UK: Cambridge University Press. ISBN 9780521707725.
Wang, W.; Lu, A.-M.; Ren, Y.; Endress, M.E. & Chen, Z.-D. (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. doi:10.1016/j.ppees.2009.01.001.
Watson, L. & Dallwitz, M.J. (1992–2006). "The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 29th July 2006". Archived from the original on 3 January 2007. Retrieved 27 December 2006.
Willemstein, S. C. (1987). An Evolutionary Basis for Pollination Ecology. E. J. Brill / Leiden University Press Leiden. ISBN 90-04-08457-6. (pbk.)

Colombo, M. L., & Bosisio, E. (1996). PHARMACOLOGICAL ACTIVITIES OFCHELIDONIUM MAJUSL. (PAPAVERACEAE). Pharmacological Research, 33(2), 127–134. https://doi.org/10.1006/phrs.1996.0019

Erdtman (1952, 1969), Faegri and Iversen (1975), Henderson (1965), Kuprianova and Alyoshina (1972, 1978), PAPAVERACEAE https://www.sciencedirect.com/sdfe/pdf/download/eid/1-s2.0-0034666779900344/first-page-pdf

Güler, D. A., Aydın, A., Koyuncu, M., Parmaksız, İ., & Tekin, Ş. (2016). Anticancer Activity of Papaver Somniferum. Journal of the Turkish Chemical Society, Section A: Chemistry, 3(3). https://doi.org/10.18596/jotcsa.43273

Labanca, F., Ovesnà, J., & Milella, L. (2018). Papaver somniferum L. taxonomy, uses and new insight in poppy alkaloid pathways. Phytochemistry Reviews, 17(4), 853–871. https://doi.org/10.1007/s11101-018-9563-3

Li, Y., Winzer, T., He, Z., & Graham, I. A. (2020). Over 100 Million Years of Enzyme Evolution Underpinning the Production of Morphine in the Papaveraceae Family of Flowering Plants. Plant Communications, 1(2), 100029. https://doi.org/10.1016/j.xplc.2020.100029

Native Plants (PAPAVERACEAE, THE POPPY FAMILY, n.d.,) https://nativeplants.org/wp-content/uploads/papaveraceae.pdf

Nemat, O. (2023). MAKING THE POPPY BAN STICK. World Today, 79(4), 19–22.