Wild carrot ( Daucus carota subsp. Carota ), illustration
The umbelliferae or umbelliferae (Apiaceae or Umbelliferae) are a family of plants in the order of the umbelliferae (Apiales). Most of the species are herbaceous plants with leaves that are divided several times and double umbels as an inflorescence , making them easy to assign to the family.
The family contains about 434 genera with about 3780 species , and is represented worldwide in the temperate zones . The umbelliferae include many aromatic plants and food plants , but also some very poisonous plant species, for example the water hemlock and the spotted hemlock .
Description and ecology
The representatives of the umbelliferae are almost exclusively perennial herbaceous plants. A few taxa, such as in the subfamily Mackinlayoideae, are lignified . The stem axis is usually hollow and nodular . The stature heights range from several meter high plants in the steppes of Central Asia ( Ferula ) to a few centimeter high cushion plants in the Antarctic ( Azorella ).
The inflorescence is usually a multi-rayed double umbel , an umbel made of usually many umbel . This structure of the inflorescences is very characteristic of the umbelliferae and has also given them their old scientific name Umbelliferae (umbrella carrier).
The bracts of the umbels are tightly packed together and form the envelope (involucre), they are often only weakly developed or absent. This is where the dome stalks originate = umbel rays. The umbellules are in turn surrounded by a (often missing) envelope (involucellum). The flower stalks are not called "dome rays". Often the inflorescence forms a dome or even an area on which insects can often be found.
Simple umbels are less common. There are also species with single flowers ( Azorella ). In the case of very large species, several double umbels can be combined to form an even larger inflorescence ( giant hogweed , Heracleum mantegazzianum ).
With the exception of the gynoeceum, the mostly inconspicuous flowers are five-fold and usually radially symmetrical . In some species, however, the marginal florets in particular are asymmetrical and therefore zygomorphic . There are originally five sepals , but they are often stunted or absent. The five petals are free and are mostly white, more rarely yellow, pink to purple. The petals often have a knocked-in lobule (lobulum inflexum) at the tip. Its shape as well as the shape of the front edge of the petal (flexure edge) are important determinants.
There is only one circle with five free, fertile stamens that are curved in the bud. Two carpels have become an under constant ovary grown. The two styluses (also known as beak) sit on a disc-shaped to conical, shiny, stylus cushion (Stylopodium). This serves as a nectarium , i.e. i.e., it excretes nectar . In each of the two ovary compartments there is a hanging anatropic ovule . A second withers away very early.
The structure of the flower can be summarized in the following flower formula :
Fruits and seeds
The fruit is a dry, two-part split fruit , also known as a double achene. The shape is often cylindrical with a round to elliptical cross-section. Spherical ( Coriandrum ) and double- spherical shapes ( Bifora ) are rarer . The two partial fruits (mericarpies or carpids) initially remain with the upper side on a fruit holder ( carpophor ), which is located in the middle.
Each partial fruit has five longitudinal ribs or main ribs (juga primaria) each with a vascular bundle on its free side. In between there are valleys (valleculae), in each of the walls of which there is a mostly dark schizogenic oil duct (here called oil streak) (caused by the divergence of cells). In some species, each valley has a secondary rib (jugum secundarium, for example the carrot with prickly secondary ribs). The oil ducts can also be increased ( Pimpinella ) or reduced ( Coriandrum ) or completely absent ( Conium ).
The seed consists of a very small embryo in a large, fat and protein-rich endosperm . The embryo lies at the top of the seed with the hypocotyl pointing upwards . The seed coat is glued to the pericarp.
The main components of the essential oils can be mainly formed from terpenes or from phenylpropanoids , depending on the type . With coriander it is predominantly (+) - linalool (terpene), with caraway (+) - carvone (terpene), with fennel and aniseed anethole (phenylpropanoid).
( S ) - (+) - carvone
Psoralen , a linear furanocoumarin
cis -Anethol (top) and trans -Anethol (bottom)
Falcarinol , a polyacetylene
The umbelliferae are the family with the largest range of coumarin compounds. In addition to simple coumarins and hydroxycoumarins (e.g. umbelliferone ), there are also a large number of prenylated, geranylated and farnesylated coumarin derivatives. This also includes the furano and pyranocoumarins. The former can be linear or angular. Hydroxy and furanocoumarins have a deterrent effect on herbivores (deterrent), as phytoalexins and as germination inhibitors. The toxicity increases from hydroxy to linear to angular furanocoumarins. The furanocoumarins are phototoxic : when exposed to UV light, the DNA is inactivated (photosensitization). Angular furanocoumarins are more toxic than linear ones, although their phototoxicity is less. Most of the species-rich genera of the family, which are widespread in the Holarctic, contain furanocoumarins ( e.g. Bupleurum and Pimpinella with 150 species each), while many monotypical genera with limited geographical distribution do not contain furanocoumarins.
Sesquiterpene lactones are found in the family with over 100 compounds. The same basic structures (e.g. germacranolide, eudesmanolide, eremophilanolide and elemanolide) occur as in the sunflower plants, but they are stereochemically different. They are also more frequently hydroxylated and esterified, especially at C11.
Over 150 polyacetylene compounds have been detected in the umbelliferous plants . The most common are the C 17 diyne dienes of the falcarinol group. The toxicity of the water hemlock ( Cicuta virosa ) and the saffron vine umbels ( Oenanthe crocata ) is based on polyacetylenes.
Alkaloids are rare. Coniin and similar piperidine derivatives occur in the spotted hemlock ( Conium maculatum ). Triterpene saponins often occur in the subfamily Saniculoideae . Typical carbohydrates are the trisaccharide umbelliferose and the sugar alcohol mannitol .
Because of the essential oils, many species are used as aromatic, vegetable and medicinal plants. The fruits, leaves and roots are used. Examples are caraway ( Carum carvi ), anise ( Pimpinella anisum ), coriander ( Coriandrum sativum ), dill ( Anethum graveolens ), lovage ( Levisticum officinale ), fennel ( Foeniculum vulgare ), parsley ( Petroselinum crispum ), and celery ( Apium graveolens ) .
Some species are very poisonous. The spotted hemlock ( Conium maculatum ) provided the poison for Socrates, who was sentenced to death . The water hemlock ( Cicuta virosa ) is also very poisonous . Dog parsley ( Aethusa cynapium ) is less poisonous , but it is often confused with parsley , which often leads to poisoning.
Many species are photosensitizing and phototoxic due to their furanocoumarins . The giant hogweed ( Heracleum mantegazzianum ) is particularly worth mentioning here . However, the psoralen contained in the phototoxic species is used in medicine as part of PUVA therapy for the treatment of skin diseases.
The family is distributed worldwide, but the main focus is in the northern temperate zones . In the tropics , umbelliferae are particularly widespread in the montane altitudes . The umbelliferae grow mainly in steppes, swamps, meadows and forests.
Synonyms for Apiaceae Lindl. nom. cons. are: Umbelliferae Juss. nom. cons., Actinotaceae AIKonstant. & Melikyan , Ammiaceae Bercht. & J.Presl , Angelicaceae Martinov , Daucaceae Martinov , Ferulaceae Sacc. , Saniculaceae Bercht. & J. Presl .
The sister taxon of the umbelliferae within the order Apiales is the group of Pittosporaceae , Araliaceae and Myodocarpaceae . The family contains about 434 genera with about 3780 species . Since 2010 it has only been divided into three subfamilies.
- The Mackinlayoideae Plunkett & Lowry are mostly woody plants. Most species grow in the areas around the South Pacific, the genus Centella mainly in South Africa . There are about six genera with about 67 species:
- Actinotus Labill. : All 15 species are common in Australia and one of them is also found in New Zealand .
- Apiopetalum Baill. : The four or so species occur in New Caledonia .
Centella L. (Syn .: Trisanthus Lour. ): The approximately 40 species are mainly distributed in the southern hemisphere , for example:
- Indian pennywort ( Centella asiatica (L.) Urb. )
- Mackinlaya F. Muell. (Syn .: Anomopanax Harms ): The ten or so species occur in Sulawesi , the Philippines , New Guinea , the Solomon Islands and northeastern Australia.
- Micropleura Lag .: There are about two species in Colombia and Chile .
- Platysace Bunge : The approximately 26 species are widespread in Australia, but none are found in Tasmania .
- Xanthosia Rudge : The 20 or so species are widespread in temperate Australia.
- The Azorelloideae Plunkett & Lowry occur in South America, Australia and the Antarctic. They have a large nucellus, a tetrasporous embryo sac. The fruits have a lignified perisperm. It contains about 21 to 23 genera with about 155 species:
- Asteriscium Cham. & Schltdl. : It contains only one type:
Andean cushion ( Azorella Lam. , Syn .: Fragosa Ruiz & Pav. , Laretia Gillies & Hook. , Mulinum Pers. ): It contains about 70 species, including:
- Yareta ( Azorella compacta Phil. )
- Bolax Comm. ex Juss. : The four to five species occur in temperate southern South America in Argentina and Chile.
- Bowlesia Ruiz & Pav .: It is mainly found in South America.
Dichosciadium Domin : It contains only one species:
- Dichosciadium ranunculaceum (F.Muell.) Domin (Syn .: Azorella ranunculacea (F.Muell.) Druce , Dichopetalum ranunculaceum F.Muell. , Azorella dichopetala Benth. Nom. Illeg., Pozoa ranunculacea (F.Muell.) Drude ): It occurs with two varieties only in the Australian states of New South Wales , Victoria and Tasmania .
- Dickinsia Franch. (Syn .: Cotylonia C. Norman ): It contains only one species:
- Diplaspis Hook. f. : The only two species occur only in the Australian states of New South Wales, Victoria and Tasmania.
- Diposis DC. : The two or three types occur in Uruguay and Chile.
- Domeykoa Phil .: The four or so species occur in Peru and Chile.
- Drusa DC. : It contains only one type:
- Eremocharis Phil .: The nine or so species occur in Peru and Chile.
- Gymnophyton Clos : The six or so species thrive in the Andes of Chile and Argentina.
- Hermas L .: There are about seven species common in southern Africa.
- Homalocarpus Hook. & Arn. : The four to six species occur in Chile.
- Huanaca Cav. : The four or so species occur in Peru and Chile.
- Oschatzia Walp. : The only two species occur only in the Australian states of New South Wales, Victoria and Tasmania.
- Pozoa Lag .: The roughly two species thrive in the Andes of Chile and Argentina.
Schizeilema (Hook. F.) Domin : It contains only one species:
- Schizeilema fragoseum (F.Muell.) Domin : It occurs only in the Australian states of New South Wales and Victoria.
Spananthe Jacq. : It contains only one type:
- Spananthe paniculata Jacq. : It occurs in the Andes.
- The Apioideae Drude , including the taxa of the former Saniculoideae Burnett . This subfamily has no stipules. The ovules are tenuinucellate, the endocarp consists of a layer of cells and is not lignified. The leaves are usually divided several times or undivided. The inflorescences are usually double umbels. The partial fruits can have a fruit holder (carpophor) and have more or less pronounced oil ducts. They have terpenoids of the Kauren-type. With the taxa of the former Saniculoideae there are now over 400 genera with around 3500 species. They are distributed worldwide, but mainly in the northern temperate zones. Lignify a few taxa, for example some Bupleurum and Myrrhidendron species. The basic chromosome number is x = 11 (8, 9, 12). For the internal systematics of this subfamily see main article: Apioideae .
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