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Apiaceae

Apiaceae (/pˈsiˌ, -sˌ/ ) or Umbelliferae is a family of mostly aromatic flowering plants named after the type genus Apium, and commonly known as the celery, carrot, or parsley family, or simply as umbellifers. It is the 16th-largest family of flowering plants, with more than 3,800 species in about 446 genera, including such well-known, and economically important plants as ajwain, angelica, anise, asafoetida, caraway, carrot, celery, chervil, coriander, cumin, dill, fennel, lovage, cow parsley, parsley, parsnip and sea holly, as well as silphium, a plant whose exact identity is unclear and which may be extinct.

The family Apiaceae includes a significant number of phototoxic species, such as giant hogweed, and a smaller number of highly poisonous species, such as poison hemlock, water hemlock, spotted cowbane, fool's parsley, and various species of water dropwort.

Most Apiaceae are annual, biennial or perennial herbs (frequently with the leaves aggregated toward the base), though a minority are woody shrubs or small trees such as Bupleurum fruticosum. Their leaves are of variable size, and alternately arranged, or with the upper leaves becoming nearly opposite. The leaves may be petiolate or sessile. There are no stipules but the petioles are frequently sheathing, and the leaves may be perfoliate. The leaf blade is usually dissected, ternate, or pinnatifid, but simple, and entire in some genera, e.g. Bupleurum. Commonly, their leaves emit a marked smell when crushed, aromatic to fetid, but absent in some species.

The defining characteristic of this family is the inflorescence, the flowers nearly always aggregated in terminal umbels, that may be simple or more commonly compound, often umbelliform cymes. The flowers are usually perfect (hermaphroditic), and actinomorphic, but there may be zygomorphic flowers at the edge of the umbel, as in carrot (Daucus carota) and coriander, with petals of unequal size, the ones pointing outward from the umbel larger than the ones pointing inward. Some are andromonoecious, polygamomonoecious, or even dioecious (as in Acronema), with a distinct calyx, and corolla, but the calyx is often highly reduced, to the point of being undetectable in many species, while the corolla can be white, yellow, pink or purple. The flowers are nearly perfectly pentamerous, with five petals and five stamens. There is often variation in the functionality of the stamens even within a single inflorescence. Some flowers are functionally staminate (where a pistil may be present but has no ovules capable of being fertilized) while others are functionally pistillate (where stamens are present but their anthers do not produce viable pollen). Pollination of one flower by the pollen of a different flower of the same plant (geitonogamy) is common. The gynoecium consists of two carpels fused into a single, bicarpellate pistil with an inferior ovary. Stylopodia support two styles, and secrete nectar, attracting pollinators like flies, mosquitoes, gnats, beetles, moths, and bees. The fruit is a schizocarp consisting of two fused carpels that separate at maturity into two mericarps, each containing a single seed. The fruits of many species are dispersed by wind but others such as those of Daucus spp., are covered in bristles, which may be hooked in sanicle Sanicula europaea and thus catch in the fur of animals. The seeds have an oily endosperm and often contain essential oils, containing aromatic compounds that are responsible for the flavour of commercially important umbelliferous seed such as anise, cumin and coriander. The shape and details of the ornamentation of the ripe fruits are important for identification to species level.

Apiaceae was first described by John Lindley in 1836. The name is derived from the type genus Apium, which was originally used by Pliny the Elder circa 50 AD for a celery-like plant. The alternative name for the family, Umbelliferae, derives from the inflorescence being generally in the form of a compound umbel. The family was one of the first to be recognized as a distinct group in Jacques Daleschamps' 1586 Historia generalis plantarum. With Robert Morison's 1672 Plantarum umbelliferarum distribution nova it became the first group of plants for which a systematic study was published.

The family is solidly placed within the Apiales order in the APG III system. It is closely related to Araliaceae and the boundaries between these families remain unclear. Traditionally groups within the family have been delimited largely based on fruit morphology, and the results from this have not been congruent with the more recent molecular phylogenetic analyses. The subfamilial and tribal classification for the family is currently in a state of flux, with many of the groups being found to be grossly paraphyletic or polyphyletic.

Prior to molecular phylogenetic studies, the family was subdivided primarily based on fruit characteristics. Molecular phylogenetic analyses from the mid-1990s onwards have shown that fruit characters evolved in parallel many times, so that using them in classification resulted in units that were not monophyletic. In 2004, it was proposed that Apiaceae should be divided into four subfamilies:

Apioideae is by far the largest subfamily with about 90% of the genera. Most subsequent studies have supported this division, although leaving some genera unplaced. A 2021 study suggested the relationships shown in the following cladogram.

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