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The families of flowering plants.                                                                                                                                                                

Umbelliferae Juss.

                       

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AlternativelyApiaceae Lindl. (sensu lato, nom. altern.)

IncludingCoriandraceae Burnett, Daucaceae Augier ex Martinov, Hydrocotylaceae (Drude) Hylander, Saniculaceae A. Löve & D. Löve

Habit and leaf form. Herbs (mostly), or shrubs (some), or ‘arborescent’, or trees (few); bearing essential oils, or without essential oils; resinous, or not resinous. Switch-plants (occasionally), or ‘normal’ plants; occasionally with the principal photosynthesizing function transferred to stems (e.g., Platysace compressa), or phyllodineous (e.g. Lilaeopsis). Leaves well developed (usually), or much reduced (dometimes, in switch forms). Plants succulent (occasionally, e.g. Crithmum), or non-succulent; autotrophic. Annual, or biennial, or perennial; with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves. Helophytic, or mesophytic, or xerophytic (e.g., Eryngium). Heterophyllous (e.g. Apium inundatum, with capillary-segmented submerged leaves), or not heterophyllous. Leaves small to large; alternate, or alternate and opposite (the upper sometimes more or less opposite); ‘herbaceous’ (usually), or leathery (occasionally), or fleshy (rarely); petiolate, or perfoliate (e.g., the upper leaves of Bupleurum rotundifolium); more or less sheathing. Leaf sheaths with free margins. Leaves gland-dotted, or not gland-dotted; aromatic, or foetid, or without marked odour (rarely); simple, or compound; peltate (sometimes), or not peltate; pulvinate, or epulvinate; when compound, ternate, or pinnate, or bipinnate, or multiply compound, or palmate (rarely). Lamina when simple, entire (commonly in Hydrocotyloideae), or dissected (usually); when simple/dissected, pinnatifid (usually), or palmatifid (e.g., in Sanicula, Astrantia,Eryngium), or spinose (e.g., in Eryngium); pinnately veined, or palmately veined, or parallel-veined. Leaves stipulate (Hydrocotyloideae), or exstipulate (usually, but sometimes with ‘stipular flanges’); without a persistent basal meristem.

Leaf anatomy. Stomata present; anomocytic, or paracytic.

Adaxial hypodermis present, or absent. Lamina dorsiventral (usually), or isobilateral, or centric; with secretory cavities. Secretory cavities containing oil, or containing resin, or containing mucilage. Minor leaf veins with phloem transfer cells (Eryngium), or without phloem transfer cells (Aegopodium, Sanicula, Smyrnium).

Stem anatomy. Stems often with hollow internodes. Secretory cavities present; with resin, or with oil, or with mucilage. Nodes multilacunar (usually), or tri-lacunar. Cortical bundles present (commonly), or absent. Medullary bundles present, or absent. Secondary thickening developing from a conventional cambial ring, or anomalous; from a single cambial ring. Xylem with libriform fibres; with vessels. Vessel end-walls simple (usually), or scalariform. Wood parenchyma paratracheal (scanty to vasicentric). Sieve-tube plastids S-type.

Reproductive type, pollination. Plants hermaphrodite (usually), or andromonoecious, or polygamomonoecious, or dioecious (Acronema). Pollination entomophilous.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in umbels (nearly always), or in heads. The ultimate inflorescence unit cymose, or racemose. Inflorescences terminal; often cymose umbels or heads arranged in cymose inflorescences, sometimes reduced to single flowers; with involucral bracts (usually, and/or with involucels), or without involucral bracts; pseudanthial (often, this phenomenon commonly associated with sterile flowers at the periphery), or not pseudanthial. Flowers bracteate; mostly small; regular to somewhat irregular. The floral irregularity involving the perianth (the corolla only). Flowers 5 merous (except for the gynoecium); cyclic; tetracyclic. Floral receptaclewith neither androphore nor gynophore. Free hypanthium absent.

Perianthwith distinct calyx and corolla (usually, but the calyx usually very reduced), or sepaline (corolla rarely absent), or petaline (calyx teeth sometimes lacking); 4–10; 2 whorled, or 1 whorled (rarely); isomerous. Calyx when detectable, 5; 1 whorled; polysepalous, or gamosepalous (often reduced to a mere rim, but never exhibiting a calyx tube); lobulate, or blunt-lobed, or toothed; persistent; with the median member posterior. Corolla 5; 1 whorled;polypetalous; valvate; unequal but not bilabiate, or regular; white, or yellow, or pink, or purple.

Androecium5. Androecial members free of the perianth; all equal to markedly unequal; free of one another; 1 whorled. Androecium exclusively of fertile stamens. Stamens 5; isomerous with the perianth; oppositisepalous; inflexed in bud. Anthers dorsifixed, or basifixed; dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Anther wall initially with one middle layer; of the ‘dicot’ type. Tapetum glandular. Pollen grains aperturate; (2–)3 aperturate; mostly (tri-) colporate; 3-celled (recorded in 16 genera).

Gynoecium 2 carpelled. The pistil 1 celled, or 2 celled. Gynoecium syncarpous; synovarious; inferior. Ovary (1–)2 locular. Gynoecium median. Epigynous disk present. Gynoecium stylate. Styles 2; free to partially joined (their bases thickened into one or two stylopodes crowning the ovary); apical. Stigmas wet type; non-papillate; Group IV type. Placentation axile, or apical. Ovules 1 per locule, or 2 per locule (usually two, but one abortive); pendulous; epitropous; with ventral raphe; non-arillate; anatropous; unitegmic; tenuinucellate, or pseudocrassinucellate. Endothelium differentiated. Embryo-sac development Polygonum-type, or Allium-type, or Penaea-type, or Drusa-type. Antipodal cells formed; 3–11; proliferating, or not proliferating. Hypostase present, or absent. Endosperm formation nuclear. Embryogeny solanad.

Fruit non-fleshy; a schizocarp. Mericarps 2; dry, united facially, 1-seeded, the integument sometimes united with the pericarp. Seeds endospermic. Endosperm oily. Embryo well differentiated (often small). Cotyledons 2. Embryo achlorophyllous (10/10); straight.

Seedling.Germination phanerocotylar.

Physiology, biochemistry. Not cyanogenic. Polyacetylenes recorded (falcarinone). Alkaloids present, or absent (but usually toxic via polyacetylenes). Iridoids not detected. Arthroquinones detected (Bupleurum, Heracleum); polyacetate derived. Proanthocyanidins absent. Flavonols present, or absent; kaempferol, or kaempferol and quercetin (mostly both). Ellagic acid absent (10 species, 10 genera). Arbutin absent. Saponins/sapogenins present, or absent. Aluminium accumulation not found. Sugars transported as sucrose (Bupleurum). Inulin not found (umbelliferose recorded). C3 (usually), or CAM (rarely). C3 physiology recorded directly in Aciphylla, Crithmum, Daucus,Eryngium, Lomatium, Pastinaca, Pituranthos, Sium. CAM recorded directly in Lilaeopsis (aquatic CAM only). Anatomy non-C4 type (Eryngium, Crithmum, Lomatium, Prangos, Sium).

Geography, cytology. Frigid zone to tropical. Cosmopolitan, but mainly North temperate. X = (4-)8–11(-12).

Taxonomy.Subclass Dicotyledonae; borderline Tenuinucelli. Dahlgren’s Superorder Araliiflorae; Araliales. Cronquist’s Subclass Rosidae; Apiales. APG 3 core angiosperms; core eudicot; Superorder Asteranae; campanulid; Order Apiales.

Species about 2850. Genera about 420; Aciphylla, Acronema, Actinolema,Actinotus, Adenosciadium, Aegopodium, Aethusa, Aframmi,Afrocarum, Afroligusticum, Afrosison, Agasyllis, Agrocharis,Ainsworthia, Albovia, Alepidea, Aletes, Alococarpum,Ammi, Ammodaucus, Ammoides, Ammoselinum, Anethum,Angelica, Anginon, Angoseseli, Anisopoda, Anisosciadium,Anisotome, Annesorhiza, Anthriscus, Aphanopleura, Apiastrum,Apium, Apodicarpum, Arctopus, Arcuatopterus, Arracia,Artedia, Asciadium, Asteriscium, Astomaea, Astrantia,Astrodaucus, Astydamia, Athamanta, Aulacospermum, Autumnalia,Azilia, Azorella, Berula, Bifora, Bilacunaria,Bolax, Bonannia, Bowlesia, Bunium, Bupleurum,Cachrys, Calyptosciandium, Capnophyllum, Carlesia, Caropsis,Carum, Caucalis, Cenolophium, Centella, Cephalopodium,Chaerophyllopsis, Chaerophyllum, Chaetosciadium, Chamaele,Chamaesciadium, Chamaesium, Chamarea, Changium, Chlaenosciadium,Choritaena, Chuanminshen, Chymsydia, Ciclospermum, Cicuta,Cnidiocarpa, Cnidium, Coaxana, Conioselinum, Conium,Conopodium, Coriandrum, Cortia, Cortiella, Cotopaxia,Coulterophytum, Crenosciadium, Crithmum, Cryptotaenia,Cuminum, Cyathoselinum, Cyclorhiza, Cymbocarpum, Cymopterus,Cynosciadium, Dactylaea, Dasispermum, Daucosma, Daucus,Dethawia, Deverra, Dichisciadium, Dickinsia, Dicyclophora,Dimorphosciadium, Diplaspis, Diplolophium, Diplotaenia,Diposis, Domeykoa, Donnellsmithia, Dorema, Dracosciadium,Drusa, Ducrosia, Dystaenia, Echinophora, Elaeoselinum,Eleutherospermum, Enantiophylla, Endressia, Eremocharis,Eremodaucus, Ergocarpon, Erigenia, Eriocycla, Eriosynaphe,Eryngium, Erythroselinum, Eurytaenia, Exoacantha, Ezosciadium,Falcaria, Fergania, Ferula, Ferulago, Foeniculum,Frommia, Froriepa, Fuernrohria, Galagania, Geocaryum,Gingidia, Glaucosciadium, Glehnia, Glia, Glochidotheca,Gongylosciadium, Grafia, Grammosciadium, Hacquetia,Halosciatrum, Haplosciadium, Haplosphaera, Harbouria,Harrysmithia, Haussknechtia, Hellenocarum, Heptaptera,Heracleum, Hermas, Heteromorpha, Hladnikia, Hohenackeria,Homalocarpus, Homalosciadium, Horstrissea, Huanaca,Hyalolaena, Hydrocotyle, Itasina, Johrenia, Kadenia,Kafirnigania, Kalakia, Kandaharia, Karatavia, Karnataka,Kedarnatha, Keraymonia, Kitagawia, Klotzschia, Komarovia,Korovinia, Korshinskia, Kosopoljanskia, Koslovia, Krasnovia,Krubera, Kundmannia, Ladyginia, Lagoecia, Laretia,Laser, Laserpitium, Lecokia, Ledebouriella, Lefebvrea,Lereschia, Levisticum, Lichtensteinia, Lignocarpa, Ligusticopsis,Ligusticum, Lilaeopsis, Limnosciadium, Lisaea, Lomatium,Lomatopodium, Magadania, Magydaris, Malabaila, Mandenovia,Marlothiella, Mastigosciadium, Mathiasella, Mediasia,Meeboldia, Melanosciadium, Melanoselinum, Merwiopsis,Meum, Micropleura, Microsciadium, Mogoltavia, Molopospermum,Monizia, Mulinum, Muretia, Musineon, Myrrhidendron,Myrrhis, Myrrhoides, Naufraga, Neocryptodiscus, Neogoezia,Neonelsonia, Neoparrya, Neoplatytaenia, Neosciadium,Niphogeton, Nirarathamnos, Nothosmyrnium, Notiosciadium,Notopterygium, Oedibasis, Oenanthe, Oligocladus, Oliveria,Olymposciadium, Oropanax, Oreocome, Oreomyrrhis, Oreonana,Oreoschimperella, Oreoxis, Orlaya, Ormopterum, Ormosciadium,Orogenia, Oschatzia, Osmorhiza, Ottoa, Oxypolis,Pachyctenium, Pachypleurum, Palimbia, Paraligusticum,Paraselinum, Parasilaus, Pastinaca, Pastinacopsis, Paulita,Pedinopetalum, Pentapeltis, Perideridia, Perissocoelium,Petagnaea, Petroedmondia, Petroselinum, Peucadanum,Phellolophium, Phlojodicarpus, Phlyctodocarpa, Physospermopsis,Physospermum, Physotrichia, Pilopleura, Pimpinella,Pinda, Platysace, Pleurospermopsis, Pleurospermum, Podistera,Polemannia, Polemanniopsis, Polylophium, Polytaenia,Polyzygus, Portenschlagiella, Pozoa, Prangos, Prionosciadium,Psammogeton, Pseudocarum, Pseudorlaya, Pseudoselinum,Pternopetalum, Pterygopleurum, Ptilimnium, Ptychotis,Pycnocycla, Pyramidoptera, Registaniella, Rhabdosciadium,Rhodosciadium, Rhopalosciadium, Rhysopterus, Ridolfia,Rouya, Rumia, Rutheopsis, Sajanella, Sanicula,Saposhnikovia, Scaligeria, Scandia, Scandix, Schizeilema,Schoenolaena, Schrenkia, Schtschurowskia, Schulzia,Schumannia, Sclerochorton, Sclerotiaria, Scrithacola,Selinum, Semenovia, Seseli, Seselopsis, Shoshonea,Silaum, Sinocarum, Sinodielsia, Sinolimprichtia, Sison,Sium, Smyrniopsis, Smyrnium, Sonderina, Soranthus,Spananthe, Spermolepis, Sphaenolobium, Sphaerosciadium,Sphallerocarpus, Sphenocarpus, Sphenosciadium, Spongiosyndesmus,Spuriodaucus, Spuriopimpinella, Stefanoffia, Steganotaenia,Stenocoelium, Stenosemis, Stenotaenia, Stewartiella,Stoibrax, Symphyoloma, Synelcoscadium, Szovitsia, Taenidia,Tamamschjania, Tauschia, Tetrataenium, Thamnosciadium,Thapsia, Thaspium, Thecocarpus, Tilingia, Tinguarra,Todaroa, Tongoloa, Tordyliopsis, Tordylium, Torilis,Tornabenea, Trachydium, Trachymene, Trachysciadium,Trachyspermum, Transcaucasia, Trepocarpus, Tricholaser,Trigonosciadium, Trinia, Trochiscanthes, Turgenia, Uldinia,Vanasushava, Vicatia, Xanthogalum, Xanthosia, Xatardia,Yabea, Zeravschania, Zizia, Zosima.

General remarks. Family review, Heywood 1971. This family exemplifies the well known difficulties in distributing certain Dicot families between Dahlgren’s Araliiflorae and Corniflorae. It is equally hard to assign them with confidence to the higher level groupings Crassinucelli and Tenuinucelli. This is interesting, given that the latter evidently represent a major divergence in the Dicot line of descent (cf.Young and Watson 1970, Chase et al. 1993). It appears on the basis of recent molecular studies (cf. APG, 1998) that Hydrocotylaceae should be transferred to Araliaceae.

Economic uses, etc. Important sources of many foodstuffs and condiments: Daucus (carrot), Pastinaca (parsnip), Apium (celery), Petroselinum (parsley), Pimpinella (anise), Carum (caraway), Anethum (dill), Anthriscus (chervil), Foeniculum (fennel), Levisticum (lovage). Ornamentals:Eryngium, Angelica, Heracleum, Trachymene etc. Some with notoriously poisonous resins or alkaloids: Cicuta, Conium (hemlocks), Aethusa (fool’s parsley).

 Illustrations:

  • Technical details: Annesorrhiza.
  • Technical details: Foeniculum, Daucus, Aethusa.
  • Technical details: Angelica, Coriandrum, Eryngium, Hydrocotyle, Scandix.
  • Actinotus helianthi: Bot. Reg. 654, 1822.
  • Aethusa cynapium (J.E. Sowerby, 1861).
  • Bunium bulbocastanum: as Carum bulbocastanum, Eng. Bot. 583 (1865).
  • Cicuta virosa (J.E. Sowerby).
  • Conium maculatum (J.E. Sowerby, 1861).
  • Coriandrum sativum: Eng. Bot. 632 (1865). 
  • Eryngium maritimum: Eng. Bot. 569 (1865).
  • Oenanthe crocata (J.E. Sowerby, 1861.
  • Oenanthe fluviatilis: Eng. Bot. 599 (1865).
  • Oenanthe aquatica (J.E. Sowerby, 1861.
  • Sanicula europaea: Eng. Bot. 568 (1865).
  • Xanthosia rotundifolia(Southern Cross, photo). 
  • British Eryngium, Hydrocotyle, Sanicula (B. Ent. compilation).
  • British Meum, Foeniculum, Silaum (B. Ent. compilation).
  • British Torilis, Anthriscus, Scandix (B. Ent. compilation).
  • British Smyrnium, Pastinaca, Daucus (B. Ent. compilation).
  • British Apium, Bupleurum (B. Ent. compilation).
  • British Sison, Pimpinella, Conopodium (B. Ent. compilation).
  • British Crithmum, Oenanthe, Aethusa (B. Ent. compilation).
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