The Aristolochiaceae () are a family, the birthwort family, of

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...

s with seven genera and about 400 known species belonging to the order

Piperales Piperales is an order of flowering plants (4,170 recognized species). It necessarily includes the family Piperaceae but other taxa have been included or disincluded variously over time. Well-known plants which may be included in this order inclu ...

. The

type genus In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal f ...

is ''

Aristolochia ''Aristolochia'' () is a large plant genus with over 500 species that is the type genus of the family Aristolochiaceae. Its members are commonly known as birthwort, pipevine or Dutchman's pipe and are widespread and occur in the most diverse clima ...

'' L.

Description

They are mostly

perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...

herbaceous plant Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition of t ...

shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...

s, or

liana A liana is a long- stemmed, woody vine that is rooted in the soil at ground level and uses trees, as well as other means of vertical support, to climb up to the canopy in search of direct sunlight. The word ''liana'' does not refer to a ta ...

s. The membranous, cordate simple

leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...

are spread out, growing alternately along the stem on leaf stalks. The margins are commonly entire. No

stipule In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole). Stipules are considered part of the anatomy of the leaf of a typical flowering plant, although in many speci ...

s are present. The bizarre

flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...

s are large to medium-sized, growing in the leaf axils. They are bilaterally or radially symmetrical.

Classification

Aristolochiaceae are

magnoliid Magnoliids (or Magnoliidae or Magnolianae) are a clade of flowering plants. With more than 10,000 species, including magnolias, nutmeg, bay laurel, cinnamon, avocado, black pepper, tulip tree and many others, it is the third-largest group of ang ...

s, a basal group of

angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...

which are not part of the large categories of

monocot Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...

s or

eudicot The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons. Traditionally they were called tricolpates or non-magnoliid dicot ...

s. As of APG IV (2016), the former families

Hydnoraceae Hydnoroideae is a subfamily of parasitic flowering plants in the order Piperales. Traditionally, and as recently as the APG III system it given family rank under the name Hydnoraceae. It is now submerged in the Aristolochiaceae. It contains two ge ...

and

Lactoridaceae ''Lactoris fernandeziana'' is a flowering shrub endemic (ecology), endemic to the cloud forest of Robinson Crusoe Island, ''Masatierra'' — Robinson Crusoe Island, of the Juan Fernández Islands archipelago of Chile. It is the only extant specie ...

are included, because exclusion would make Aristolochiaceae in the traditional sense

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

. Some newer classification schemes, such as the update of the

Angiosperm Phylogeny Group The Angiosperm Phylogeny Group (APG) is an informal international group of systematic botanists who collaborate to establish a consensus on the taxonomy of flowering plants (angiosperms) that reflects new knowledge about plant relationships disco ...

, place the family Aristolochiaceae in the order Piperales, but it is still quite common, though superseded, for the Aristolochiaceae to be assigned, sometimes with some other families, their own order ( Aristolochiales).

Phylogeny

Four assemblages can be distinguished in the genus-level

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

of Aristolochiaceae: * ''Aristolochia'' is closely related to ''Thottea''. * ''Hydnora'' is closely related to ''Prosopanche''. * ''Lactoris'' occupies an isolated position. * ''Asarum'' is closely related to ''Saruma'', and both genera display a deep-branching position in the family.

Phytochemistry

Many members of ''

'' and some of ''

Asarum ''Asarum'' is a genus of plants in the birthwort family Aristolochiaceae, commonly known as wild ginger. ''Asarum'' is the genitive plural of the Latin ''āsa'' (an alternate form of ''āra'') meaning altar or sanctuary. Description ''Asarum' ...

'' contain the toxin aristolochic acid, which discourages herbivores and is known to be

carcinogen A carcinogen is any substance, radionuclide, or radiation that promotes carcinogenesis (the formation of cancer). This may be due to the ability to damage the genome or to the disruption of cellular metabolic processes. Several radioactive substan ...

ic in rats. ''Aristolochia'' species are carcinogenic to

human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...

Genomics

The complete

plastid genome Chloroplast DNA (cpDNA) is the DNA located in chloroplasts, which are photosynthetic organelles located within the cells of some eukaryotic organisms. Chloroplasts, like other types of plastid, contain a genome separate from that in the cell nuc ...

sequence of one species of Aristolochiaceae, ''

Hydnora visseri ''Hydnora visseri'', the Visser's hydnora, is a subterranean holoparasitic plant, lacking leaves and roots, and is described from southwestern Namibia and northwestern South Africa and has the longest tepal lobes of all ''Hydnora'' species. The g ...

'', has been determined. As compared to the

chloroplast genome Chloroplast DNA (cpDNA) is the DNA located in chloroplasts, which are photosynthetic organelles located within the cells of some eukaryotic organisms. Chloroplasts, like other types of plastid, contain a genome separate from that in the cell Cel ...

of its closest photosynthetic relatives, the

plastome Chloroplast DNA (cpDNA) is the DNA located in chloroplasts, which are photosynthetic organelles located within the cells of some eukaryotic organisms. Chloroplasts, like other types of plastid, contain a genome separate from that in the cell n ...

of ''Hydnora visseri'' shows extreme reduction in both size (ca. 27 kilo

base pairs A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...

) and gene content (24 genes appear to be functional). This Aristolochiaceae species therefore possesses one of the smallest plastid genomes among

Ecology

Pipevine swallowtail ''Battus philenor'', the pipevine swallowtail or blue swallowtail, Retrieved April 19, 2018. is a swallowtail butterfly found in North America and Central America. This butterfly is black with iridescent-blue hindwings. They are found in many di ...

butterflies lay their

egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...

s on pipevine (''Aristolochia'' species), and the

larvae A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...

feed on the plant, but are not affected by the

toxin A toxin is a naturally occurring organic poison produced by metabolic activities of living cells or organisms. Toxins occur especially as a protein or conjugated protein. The term toxin was first used by organic chemist Ludwig Brieger (1849– ...

, which then offers the adult butterfly protection against

predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...

Fossil record

The oldest

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

leaf remains are of †''Aristolochites dentata'' from the

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...

Nebraska Nebraska () is a state in the Midwestern region of the United States. It is bordered by South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Kansas to the south; Colorado to the southwe ...

United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...

Pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...

record of †''Aristolochiacidites viluiensis'' has been described from

Upper Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...

sediments of

Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part of ...

Fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

wood is known from the

Deccan Traps The Deccan Traps is a large igneous province of west-central India (17–24°N, 73–74°E). It is one of the largest volcanic features on Earth, taking the form of a large shield volcano. It consists of numerous layers of solidified flood ...

India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...

some 66 million years ago. Leaf fossils of ''

'' are known from the Early and Late

Tertiary Tertiary ( ) is a widely used but obsolete term for the geologic period from 66 million to 2.6 million years ago. The period began with the demise of the non-avian dinosaurs in the Cretaceous–Paleogene extinction event, at the start ...

North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...

and the Late

Abkhazia Abkhazia, ka, აფხაზეთი, tr, , xmf, აბჟუა, abzhua, or ( or ), officially the Republic of Abkhazia, is a partially recognised state in the South Caucasus, recognised by most countries as part of Georgia, which vi ...

Ukrainia Ukraine ( uk, Україна, Ukraïna, ) is a country in Eastern Europe. It is the second-largest European country after Russia, which it borders to the east and northeast. Ukraine covers approximately . Prior to the ongoing Russian invas ...

and

Poland Poland, officially the Republic of Poland, is a country in Central Europe. It is divided into 16 administrative provinces called voivodeships, covering an area of . Poland has a population of over 38 million and is the fifth-most populous ...

. Fossil leaf remains of †''Aristolochia austriaca'' have been described from

Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...

sediments Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sand a ...

of the Pellendorf site at the

Vienna Basin The Vienna Basin (german: Wiener Becken, cz, Vídeňská pánev, sk, Viedenská kotlina, Hungarian: ''Bécsi-medence'') is a geologically young tectonic burial basin and sedimentary basin in the seam area between the Alps, the Carpathians and t ...

Austria Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populous ...

. †''A. austriaca'' is most similar to the extant

Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the e ...

species ''A. rotunda'' and ''A. baetica''.The first fossil Aristolochia (Aristolochiaceae, Piperales) leaves from Austria by Barbara Meller, Article number: 17.2.21A, https://doi.org/10.26879/420, Palaeontological Association, May 2014

References

External links

International Agency for Research on Cancer evaluation

U.S. Food and Drug Administration alerts

* ttp://www.botanical-dermatology-database.info/BotDermFolder/ARIS.html Aristolochiaceaei
BoDD – Botanical Dermatology Database
{{Taxonbar, from=Q156446 Magnoliid families