The Families of Angiosperms

L. Watson and M.J. Dallwitz

Umbelliferae Juss.

Alternatively Apiaceae Lindl. (sensu lato, nom. altern.).

Including Coriandraceae 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 (sometimes, in switch forms). Plants succulent (occasionally, e.g. Crithmum), or non-succulent. Annual, or biennial, or perennial; with a basal aggregation of leaves, or without conspicuous aggregations of leaves. Helophytic, or mesophytic, or xerophytic (e.g., Eryngium). Conspicuously heterophyllous (e.g. Apium inundatum, with capillary-segmented submerged leaves), or not conspicuously 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); when pinnate, imparipinnate. 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’); leaf development not ‘graminaceous’.

Leaf anatomy. The leaf lamina dorsiventral (usually), or bifacial, or centric. Stomata present; anomocytic, or paracytic. Hairs present; mostly eglandular (including unicellular, dendroid and stellate forms). Adaxial hypodermis present, or absent. Lamina with secretory cavities. Secretory cavities containing oil, containing mucilage, and containing resin (cf. those in the axes, the contents often mixed). The mesophyll not containing mucilage cells (secretory cells of any kind seemingly lacking); containing crystals, or without crystals (often). The crystals when present, generally druses. Minor leaf veins with phloem transfer cells (Eryngium), or without phloem transfer cells (Aegopodium, Sanicula, Smyrnium).

Axial (stem, wood) anatomy. Young stems often with hollow internodes. Secretory cavities present; with oil, with resin, and with mucilage (as in the leaves, containing clear, turbid, milky white or yellow mixtures, in cortex, pericycle and phloem). Cork cambium present, or absent; when present, initially deep-seated, or initially superficial. Nodes multilacunar (usually), or tri-lacunar. Primary vascular tissues principally comprising a ring of bundles; collateral. Cortical bundles present (commonly), or absent. Medullary bundles present (but rarely), or absent. Secondary thickening absent, or developing from a conventional cambial ring (?), or anomalous (occasionaly, exemplified in some species of Eryngium). The anomalous secondary thickening in Eryngium via concentric cambia (involving an extra-fascicular cambial ring), or from a single cambial ring.

The wood diffuse porous. The vessels very small to medium. The vessel end-walls simple (usually), or reticulately perforated and scalariform. The vessels without spiral thickening. The axial xylem without fibre tracheids; with libriform fibres; seemingly without septate fibres (contrasting with their widespread occurrence in Araliaceae). The fibres without spiral thickening. The parenchyma paratracheal (scanty to vasicentric); wood not storied.

Reproductive type, pollination. Unisexual flowers present, or absent. 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 units 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. Free hypanthium absent.

Perianth with 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.

Androecium 5. 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 usually abortive - cf. Araliaceae); 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, phytochemistry. 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). Sugars transported as sucrose (Bupleurum). Inulin not found (umbelliferose recorded). Not cyanogenic. Polyacetylenes recorded (falcarinone). Alkaloids present, or absent (poisonous umbellifers usually toxic via polyacetylenes). Anthraquinones detected (Bupleurum, Heracleum); polyacetate derived. Arbutin absent. Iridoids not detected. Saponins/sapogenins present, or absent. Proanthocyanidins absent. Flavonols present, or absent; kaempferol, or kaempferol and quercetin (mostly both). Ellagic acid absent (10 species, 10 genera). Aluminium accumulation not found. Sieve-tube plastids S-type.

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 III core angiosperms; core eudicot; Superorder Asteranae; campanulid. APG IV 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 reviewed by Heywood 1971. Apiales exemplify the well known difficulties in distributing certain Dicot families between Dahlgren’s Araliiflorae and Corniflorae. It was equally hard to assign them with confidence to the higher level groupings Crassinucelli and Tenuinucelli, despite the fact that the latter evidently represent a major divergence in the Dicot line of descent (cf.Young and Watson 1970, Chase et al. 1993). In terms of the data compiled for this package, other than two ‘esoteric characters’ reflecting limited sampling (wet/dry stigmas, occurrence of septate wood fibres), there are no absolute differences between Araliaceae and Umbelliferae: they are distinguishable only in having the states of overlapping characters expressed in very different proportions. This does not mean of course that the families should be merged, because the names permit useful predictive generalization, On the other hand, it does support S.M. Walters’s (1960) entertaining contention regarding the historical origins of useful family circumscriptions; viz., that if formal taxonomy had originated in the southern hemisphere, the Umbelliferae would now be a subfamily of the Araliaceae - which accords with the recent notion that Hydrocotyloideae should be transferred to the Araliaceae (cf. molecular studies and APG 3.).

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).

Quotations.

I cannot tarry: I know a wench married in an afternoon as she went to the garden for parsley to stuff a rabbit
(‘Taming of the Shrew’, v., 4)

I, with my long nails, will dig thee pig-nuts
(‘Tempest’, ii., 2 - Conopodium majus)

Let the sky . . . .
Hail kissing-comfits and snow eringoes
(‘Merry Wives’, v., 5 - candied Eryngium roots)

. . . . Half-way down
Hangs one that gathers samphire; dreadful trade!
(‘King Lear’, iv., 6 — Crithmum maritimum, formerly collected from maritime cliffs for salad and pickling)

(Poins) . . . plays at quoits well and eats conger and fennel
(‘2nd Henry the Fourth’, ii., 4 - of Foeniculum (with eel). cf. Turner, 1568, vol.2 : ‘Auttours wryte that serpentes waxe yonge agayne by tastinge and eatinge this herbe, wherefore some thynke that (its use) is very mete for aged folke’. Also, according to Ophelia, ‘It hath a wonderful propertie to take away the film or web that overcasteth and dimmeth our eyes’)

The Conium there, her stalks bedropp’d with red,
Rears, with Circaea, neighbour of the dead
(Charlotte Smith, quoted by Ann Pratt, ‘Wild Flowers’ 1857)

We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, genera included in each family, and classifications (Dahlgren; Dahlgren, Clifford, and Yeo; Cronquist; APG). See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 25th April 2024. delta-intkey.com’.

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