DIPSAPIIDAE D. C. Tank et M. J. Donoghue

Tank et Donoghue in Syst. Bot. 35(2): 434. 2010


[Araliales+Dipsidae]


ARALIALES Juss. ex Bercht. et J. Presl

Berchtold et Presl, Přir. Rostlin: 258. Jan-Apr 1820 [‘Araliae’]

Apiales Nakai, Hisi-Shokubutsu: 58. 1930; Aralianae Takht., Sist. Filog. Cvetk. Rast.: 357. 4 Feb 1967

Habit Usually bisexual (sometimes monoecious, andromonoecious, polygamomonoecious, gynomonoecious, dioecious or gynodioecious), evergreen or deciduous trees, shrubs or lianas, suffrutices, perennial, biennial or annual herbs. Some representatives are xerophytes, helophytes or aquatic. Internodes often hollow. Leaf scars often large and distinct.

Vegetative anatomy Phellogen ab initio superficially or deeply seated. Peripheral collenchyma usually well developed. Medulla large and wide (as old shrinking and disappearing). Medullary and/or cortical vascular bundles frequent, often inverted. Secondary lateral growth normal or anomalous (via cylindrical cambium). Vessel elements often solitary; often at least some vessels in groups. Vessel elements with simple or scalariform (sometimes reticulate etc.) perforation plates; lateral pits alternate, scalariform or opposite, usually bordered pits. Vestured pits often present. Imperforate tracheary xylem elements usually libriform fibres (sometimes fibre tracheids or tracheids) with usually bordered pits (often reduced bordered pits), septate or non-septate (sometimes also vasicentric tracheids). Wood rays uniseriate or multiseriate, usually heterocellular. Axial parenchyma usually paratracheal scanty vasicentric or banded (sometimes apotracheal diffuse or diffuse-in-aggregates), or absent. Tyloses sometimes frequent. Palisade mesophyll often with arm cells. Sieve tube plastids Ss type. Nodes usually ≥5:≥5, multilacunar with five or more leaf traces (sometimes 1:3 or 3:3, unilacunar or trilacunar with three traces). Parenchyma and other tissues usually with schizogenous secretory canals and cavities containing ethereal oils, resins, gums, or mucilage. Prismatic calciumoxalate crystals or crystal sand sometimes frequent.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple or branched, often vesicular, sometimes dendritic, stellate, T-shaped, peltate or lepidote; glandular hairs multicellular uniseriate, or absent.

Leaves Usually alternate (spiral or rarely distichous; rarely opposite or verticillate), simple or compound (often several times pinnately compound), usually pinnately lobed (sometimes palmately lobed or entire), with conduplicate, curved or supervolute ptyxis (rarely linear, terete, articulated leaves with hydathodes). Stipules absent or cauline (rarely intrapetiolar); lower part of petiole usually sheathingly expanding and enclosing stem. Petiole vascular bundles with very various and complex anatomy. Venation pinnate or palmate (rarely parallelodromous). Stomata usually anomocytic or paracytic (sometimes anisocytic, parallelocytic, cyclocytic or diacytic). Cuticular wax crystalloids as rodlets (Pittosporaceae). Domatia rarely present. Secretory canals and cavities with ethereal oils, resins, gums, or mucilage. Epidermis sometimes with mucilage cells. Mesophyll sometimes with sclerenchymatous idioblasts. Mesophyll cells sometimes with calciumoxalate druses. Leaf margin and leaflet margins usually serrate (rarely entire, crenate or sinuate).

Inflorescence Terminal or axillary, panicle, thyrsoid, corymb, or simple or compound umbels (of dichasial origin), or capitate (sometimes with whorls of flowers below terminal umbel), rarely racemose (flowers sometimes solitary). Floral prophylls (bracteoles, involucel bracts) subtending flowers of partial umbels, or absent. Bracts (involucral bracts) subtending partial inflorescences (partial umbels), or absent. Inflorescences sometimes pseudanthia with petaloid involucral bracts (and sometimes sterile or female peripheral flowers).

Flowers Usually actinomorphic (marginal flowers sometimes zygomorphic), often small. Pedicel sometimes articulated. Usually epigyny (sometimes hypogyny or half epigyny). Sepals five, usually minute (sometimes absent), usually with open (sometimes imbricate, rarely valvate) aestivation, persistent or caducous, free or more or less connate; median sepal adaxial. Petals (three to) five (rarely ten or twelve), usually with valvate (sometimes imbricate) aestivation, often clawed, often caducous, usually (secondarily) free (sometimes connate at base, rarely absent), with apex often inflexed. Nectariferous disc intrastaminal, annular to angular, usually present on stylopodium (nectary and disc rarely absent).

Androecium Stamens (three to) five to ten (to twelve, rarely numerous), usually haplostemonous, antesepalous, alternipetalous (rarely diplostemonous), often unequal in length. Filaments often inflexed in bud, free from each other, from petals and from pistil, inserted on or outside annular nectariferous disc. Anthers basifixed or dorsifixed, versatile or non-versatile, usually tetrasporangiate (rarely octosporangiate), usually introrse (rarely latrorse), usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–6)-colporate or (2–)3(–6)-colpate, shed as monads, usually tricellular (rarely bicellular) at dispersal. Exine tectate or semitectate, with columellate infratectum, reticulate, perforate, scabrate or striate (sometimes imperforate), with various supratectal processes or smooth.

Gynoecium Pistil composed of two (to five; rarely numerous) connate carpels (sometimes three carpels, with abaxial carpel fertile); vascular bundles of ventral carpels usually consisting of connate bundles from common placenta; when fused ventral bundles opposite carpels, then two bundles arising from common carpel; when fused ventral bundles present in septal radia, then two bundles arising from adjacent carpel. Ovary usually inferior (sometimes superior or semi-inferior), usually bilocular to quinquelocular (rarely unilocular, multilocular or pseudomonomerous). Stylodia two (to five; style sometimes single), short, usually free (rarely connate at base), usually inserted on epigynous stylopodium (on top of ovary). Stigmas capitate to punctate (rarely somewhat lobate), papillate or non-papillate, usually Wet (sometimes Dry) type. Pistillodium usually absent.

Ovules Placentation usually axile to apical (occasionally parietal). Ovules usually one (sometimes two, rarely numerous) per carpel, anatropous, pendulous (rarely horizontal or ascending), usually epitropous (rarely apotropous), unitegmic, tenuinucellar or crassinucellar (sometimes pseudocrassinucellar). Hypostase present or absent. Funicular obturator rarely present. Nucellar cap present or absent. Megagametophyte usually monosporous, Polygonum type (rarely Oenothera type, or disporous, Allium type, or tetrasporous, Penaea, or Drusa types). Antipodal cells often proliferating. Endosperm development ab initio nuclear. Endosperm haustoria absent. Embryogenesis solanad, onagrad, asterad or chenopodiad.

Fruit Usually a drupe (often dry) or dry schizocarp with persistent carpophore (formed by ventral vascular bundles supporting carpels and surrounding tissues) and two nutlike mericarps (sometimes a berry, a loculicidal capsule or a syncarp). Vallecular and intrajugal vittae, often with elongate secretory oil canals and oil cavities.

Seeds Aril usually absent. Exotestal cell walls thick or thin, often tangentially elongate, tanniniferous. Endotesta unspecialized. Perisperm not developed. Endosperm copious, oily (sometimes ruminate), with petroselinic acid. Embryo usually small, straight, well differentiated, without chlorophyll. Cotyledons (one or) two. Germination phanerocotylar.

Cytology x = (4–)8–12 (or more); x = 6? for Araliales (Yi & al. 2004)

DNA Mitochondrial gene rpl2 absent (lost). Deletion of 92 bp in plastid gene rpl16.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin?), flavones, sulfated or O-methylated flavonoids, Route I carbocyclic iridoids (griselinoside, aralioside, verbascosides) and Route II decarboxylated iridoids (rare), simple coumarins (e.g. umbelliferone), hydroxycoumarins, pyranocoumarins, dihydropyranocoumarins, furanocoumarins, dihydrofuranocoumarins etc., oleanolic acid derivatives, dammaranes, sesquiterpene lactones, triterpenoid ethereal oils and resins, caffeic acid, chlorogenic acid, ellagic acid, tannins (rare), proanthocyanidins (prodelphinidins), alkaloids (incl. benzylisoquinoline and hemlock alkaloids), triterpene saponins, acetate-derived arthroquinones, phenantrenes, phenylpropenes (e.g. myristicin), acetophenones, germacrane-like compounds, asarone, syringaresinol, pinoresinol, polyacetylenes (e.g. falcarinone) derived from fatty acids, mostly aliphatic C17 acetylenes (substitute for iridoids?), petroselinic acid (in endosperm), and mannitol. Cyanogenic compounds not found. Carbohydrates stored as trisaccharide umbelliferose.

Systematics Araliales are sister-group to Dipsidae, i.e the clade [Paracryphiaceae+Dipsacales].

A well supported topology of Araliales is the following: [Pennantiaceae+[Torricelliaceae+ [Griseliniaceae+[Pittosporaceae+[Araliaceae+[Myodocarpaceae+Apiaceae]]]]]].

Pennantia has hypogynous flowers. The pistil is composed of three connate carpels, although it is seemingly consisting of a single carpel and often only the abaxial carpel is fertile. The anatomy, pollen morphology, embryology and phytochemistry are very poorly known in Pennantia, which makes the character polarization difficult in Araliales.

Potential synapomorphies for the Araliales with the exception of Pennantia include (Stevens 2001 onwards): young stems with peripheral collenchyma; pericyclic fibres few or absent; vessel elements mainly occurring in groups and with simple perforations; pits usually simple or only narrowly bordered; paratracheal axial parenchyma scanty; nodes ≥5:≥5; leaf base broad; epigyny; petals secondarily free, with imbricate aestivation; nectary inserted on top of ovary; styles/stigmas recurved; and polyacetylenes present.

Griselinia and Torricelliaceae share many features. Both groups have vessel elements arranged in clusters; septate fibres with simple pits; wood rays sometimes more than ten cell layers wide and with square or upright cells; ovary with transseptal vascular bundles; only the abaxial carpel fertile; ovule apotropous; and presence of griselinoside. Since these characters are virtually unknown in Pennantia, it cannot be established whether they are apomorphies or plesiomorphies in Araliales.

The clade [Griseliniaceae+[Pittosporaceae+[Araliaceae+[Myodocarpaceae+Apiaceae]]]] has the potential synapomorphies: glandular hairs rare; petals single-veined; nectaries present; ovule with endothelium; and petroselinic acid present in seeds.

The clade [Pittosporaceae+[Araliaceae+[Myodocarpaceae+Apiaceae]]] has a large number of characters in common (Stevens 2001 onwards): lateral roots originating from either side of xylem poles (one resinous canal running down stem in pole apex); parenchyma in secondary phloem surrounding secretory canals and arranged in groups with sieve tubes; schizogenous resinous canals present in pericycle, phloem etc.; nodes sometimes 3:3; shoot with reduced bract-like leaves at base; leaves with conduplicate ptyxis; leaf margins serrate or otherwise incised; leaf teeth often with broad glandular apex with one main and two accessory veins, or one vein proceeding above tooth; ultimate inflorescence units umbels; corolla tube formation early; pollen grains tricellular at dispersal; carpels two, connate, both fertile, with nectariferous stylopodium delimited from style by basal furrow; stigma Wet type; ovules two per carpel, epitropous; seeds often with hemicellulose as carbohydrate reserve; embryo tiny; x = 12; presence of furanocoumarins, triterpenoid ethereal oils, falcarinone, polyacetylenes (mainly aliphatic, including C17 acetylenes, etc.), and acetate-derived anthraquinones; and absence of iridoids, flavonols and tannins.

The [Araliaceae+[Myodocarpaceae+Apiaceae]] clade is supported by, i.a., the following characters: inflorescence terminal; epigyny; calyx minute, with open aestivation; presence of stylopodium; nectary (on stylopodium) divided; one of two ovules much reduced; fruit a schizocarp; fruit laterally compressed; wings or “pseudo-wings”, when present, with mesocarp and endocarp, with vascular bundles at wing margin; deletion comprising 92 bp in plastid gene rpl16; petroselinic acid and polyacetylene C18 tariric fatty acid present in seeds; and trisaccharide umbelliferose as carbohydrate reserve.

Finally, Myodocarpaceae and Apiaceae both have inflorescence panicle or raceme; and furanocoumarins.

The floral “disc” in Apiaceae and Araliaceae, and probably in Myodocarpaceae, is in reality a carpellary margin nectary displaced due to intercalary growth. The axile placentation in these three clades may be equivalent to the parietal placentation in Pittosporaceae, in which there is a short basal zone of separate locules, the ovules being inserted above this zone. The ovules are principally inserted on corresponding sites in Apiaceae and Araliaceae.

Phylogeny of Araliales based on DNA sequence data (Tank & Donoghue 2010).

APIACEAE Lindl.

( Back to Araliales )

Lindley, Intr. Nat. Syst. Bot., ed. 2: 21. Jul 1836, nom. cons.

Umbelliferae Juss., Gen. Plant.: 218. 4 Aug 1789, nom. cons. et nom. alt.; Ammiaceae Bercht. et J. Presl, Přir. Rostlin: 259. Jan-Apr 1820; Angelicaceae Martinov, Tekhno-Bot. Slovar: 29. 3 Aug 1820 [’Angeliceae’]; Bupleuraceae Bercht. et J. Presl, Přir. Rostlin: 258. Jan-Apr 1820 [’Bupleureae’]; Caucalidaceae Bercht. et J. Presl, Přir. Rostlin: 259. Jan-Apr 1820 [‘Caucalideae’]; Daucaceae Augier ex Martinov, Tekhno-Bot. Slovar: 183. 3 Aug 1820 [’Daucoideae’]; Eryngiaceae Bercht. et J. Presl, Přir. Rostlin: 259. Jan-Apr 1820 [‘Eryngieae’]; Imperatoriaceae Martinov, Tekhno-Bot. Slovar: 328. 3 Aug 1820 [‘Imperatoriae’]; Lagoeciaceae Bercht. et J. Presl, Přir. Rostlin: 259. Jan-Apr 1820 [‘Lagoeciae’]; Pastinacaceae Martinov, Tekhno-Bot. Slovar: 457. 3 Aug 1820 [‘Pastinaceae’]; Pimpinellaceae Bercht. et J. Presl, Přir. Rostlin: 258. Jan-Apr 1820 [‘Pimpinelleae’]; Saniculaceae Bercht. et J. Presl, Přir. Rostlin: 259. Jan-Apr 1820 [’Saniculeae’]; Scandicaceae Bercht. et J. Presl, Přir. Rostlin: 259. Jan-Apr 1820 [’Scandicinae’]; Selinaceae Bercht. et J. Presl, Přir. Rostlin: 259. Jan-Apr 1820 [’Selineae’]; Sileraceae Bercht. et J. Presl, Přir. Rostlin: 259. Jan-Apr 1820; Ammiineae Link, Handbuch 1: 327. 4-11 Jul 1829 [‘Ammieae’]; Bupleurineae Link, Handbuch 1: 317. 4-11 Jul 1829 [‘Bupleurinae’]; Pimpinellineae Link, Handbuch 1: 506. 4-11 Jul 1829 [‘Pimpinelleae’]; Angelicales Burnett, Outl. Bot.: 1127. Jun 1835 [‘Angelicinae’]; Coriandraceae Burnett, Outl. Bot.: 773, 783, 1093, 1128. Feb 1835; Smyrniaceae Burnett, Outl. Bot.: 773, 1093, 1128. Feb 1835; Angelicineae Burnett, Outlines Bot.: 616, 762, 1093, 1127. Feb 1835 [‘Angelicosae’]; Caucalidineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 702, 753. 1846 [‘Caucalideae’]; Coriandrineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 702, 766. 1846 [‘Coriandreae’]; Daucineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 702, 752. 1846; Saniculineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 701, 707. 1846 [‘Saniculeae’]; Scandicineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 702, 755. 1846 [‘Scandicinae’]; Silerineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 701, 748. 1846; Smyrniineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 702, 759. 1846 [‘Smyrneae’]; Ferulaceae Sacc. in Nuovo Giorn. Bot. Ital. 4: 214. 31 Jul 1872; Ammiales Small, Fl. S.E. U.S.: 851. 22 Jul 1903; Mackinlayaceae Doweld, Tent. Syst. Plant. Vasc.: lii. 23 Dec 2001; Apiineae G. M. Plunkett et Lowry in G. M. Plunkett et al., S. African J. Bot. 70: 379. 7 Oct 2004; Actinotaceae Konstant. et Melikjan in Bot. Žurn. 90: 1763. 15 Nov 2005

Genera/species 450–460/3.460–3.580

Distribution Cosmopolitan except polar areas.

Fossils Fossils of Apiaceae are reported from some Cenozoic sites.

Habit Usually bisexual (sometimes andromonoecious or polygamomonoecious, rarely dioecious), usually perennial, biennial or annual herbs (sometimes suffrutices or shrubs, rarely trees). Some representatives are xerophytic, aquatic or helophytic. Internodes usually hollow.

Vegetative anatomy Phellogen ab initio superficially or cortical. Peripheral collenchyma usually well developed. Medulla large and wide (shrinking and disappearing as old). Medullary and cortical vascular bundles abundant. Secondary lateral growth normal or anomalous (from cylindrical cambium). Vessel elements usually with simple (sometimes scalariform; in Mackinlayoideae also reticulate or foraminate) perforation plates; lateral pits alternate, scalariform or opposite, bordered pits. Vestured pits? Imperforate tracheary xylem elements libriform fibres with simple or (reduced) bordered pits, septate or non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma usually paratracheal scanty vasicentric or banded (rarely apotracheal diffuse or diffuse-in-aggregates). Sieve tube plastids S type. Nodes usually ≥5:≥5, multilacunar with five or more leaf traces (sometimes 3:3, trilacunar with three traces). Parenchyma and other tissues usually with schizogenous secretory canals and cavities containing ethereal oils, resins, gums, or mucilage. Druses sometimes frequent. Prismatic calciumoxalate crystals sometimes frequent (in, e.g., Mackinlayoideae).

Trichomes Hairs unicellular or multicellular, often vesicular, uniseriate or branched, sometimes dendritic or stellate; glandular hairs frequent.

Leaves Usually alternate (spiral; rarely opposite or verticillate), simple or compound (often several times pinnately compound), usually pinnately lobed (sometimes palmately lobed or entire; rarely linear-terete and articulated with hydathodes at articulations), with ? or supervolute ptyxis. Stipules usually absent (present in some genera in Azorelloideae); lower part of petiole usually sheathingly expanding and enclosing stem (leaf sheath often long). Petiole vascular bundles with very various and complex anatomy. Venation usually pinnate (sometimes palmate, rarely parallelodromous). Stomata usually anomocytic or paracytic (sometimes anisocytic, parallelocytic, or diacytic). Cuticular wax crystalloids as platelets, irregular structures, etc. Secretory canals and cavities with ethereal oils, resins, gums, or mucilage. Leaf margin and leaflet margins, respectively, usually serrate (rarely entire); each tooth with broad glandular apex with one main vein and two lateral veins, or with one vein running above tooth.

Inflorescence Usually terminal (sometimes axillary), simple or usually compound umbels or capitate (occasionally with whorls of flowers below terminal umbel; flowers sometimes solitary). Floral prophylls (bracteoles) subtending flowers of partial umbels, or absent. Bracts subtending partial inflorescences (partial umbels), or absent. Inflorescences sometimes pseudanthia with petaloid involucral bracts (and possibly sterile peripheral flowers).

Flowers Usually actinomorphic (marginal flowers often zygomorphic), small. Epigyny. Sepals usually minute or entirely reduced (in Saniculoideae five, larger), with valvate or open aestivation, persistent, free or more or less connate; median sepal adaxial. Petals usually five (rarely four), with valvate aestivation, usually clawed, free (rarely absent), with apex usually inflexed. Nectariferous disc intrastaminal, annular, usually on stylopodium.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous, often unequal in length. Filaments inflexed in bud, free from each other and from petals, inserted on nectariferous disc. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (rarely dicolporate), shed as monads, tricellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate, reticulate, or striate, with various supratectal processes or smooth.

Gynoecium Pistil composed of two connate carpels; ventral carpel vascular bundles consisting of connate bundles from common placenta. Ovary inferior, usually bilocular (rarely unilocular, pseudomonomerous). Stylodia two, short, free, usually inserted on stylopodium on top of ovary. Stigmas capitate to punctate, non-papillate, Wet type. Pistillodium?

Ovules Placentation axile to apical. Ovules one or two (one fertile and one sterile) per carpel, anatropous, pendulous, epitropous, unitegmic, usually tenuinucellar (sometimes pseudocrassinucellar). Integument approx. seven cell layers thick. Nucellar cap present or absent. Hypostase present or absent. Megagametophyte usually monosporous, Polygonum type (rarely disporous or tetrasporous, Oenothera, Allium, Penaea, or Drusa type). Antipodal cells often proliferating. Endosperm development ab initio nuclear. Endosperm haustoria absent. Embryogenesis solanad.

Fruit Usually a dry schizocarp with persistent carpophore (formed from ventral vascular bundles supporting carpels and surrounding tissues; rarely absent) and two nutlike mericarps, often with four or five ridges or wings, and furrows (fruit rarely fleshy). Mesocarp usually lignified. Vallecular and intrajugal vittae with elongate secretory oil canals and oil cavities usually present. Crystals solitary, rhomboidal, in small-celled inner layer. Endocarp usually lignified, two cell layers thick, sclereidal-fibrous. Testa sometimes adnate to endocarp.

Seeds Aril absent. Exotestal cells thin-walled. Endotesta unspecialized. Perisperm not developed. Endosperm copious, oily, with petroselinic acid. Embryo usually small, straight, well differentiated, without chlorophyll. Cotyledons usually two (rarely one). Germination phanerocotylar.

Cytology x = (4–)8–12 (13 or more)

DNA Mitochondrial gene rpl2 absent. Deletion of 92 bp present in plastid gene rpl16. Plastid inverted repeat in Coriandrum comprising less than 12 kb.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin?), sulphated or O-methylated flavonoids (in Apioideae), simple coumarins (e.g. umbelliferone), pyranocoumarins, dihydropyranocoumarins, furanocoumarins, dihydrofuranocoumarins etc., sesquiterpene lactones, triterpenoid ethereal oils and resins, caffeic acid, hemlock alkaloids, benzylisoquinoline alkaloids, triterpene saponins, polyols, acetate-derived arthroquinones, phenylpropenes (e.g. myristicin), polyacetylenes (e.g. falcarinone) derived from fatty acids, mostly aliphatic C17 acetylenes, petroselinic acid (in endosperm), lignans, and mannitol present. Flavones, hydroxycoumarins, ellagic acid, tannins, proanthocyanidins and cyanogenic compounds not found. Carbohydrates stored as umbelliferose (trisaccharide, raffinose isomer).

Use Ornamental plants, spices, flavours, perfumes, vegetables, medicinal plants, poisons, gum resins (Ferula spp.).

Systematics Apiaceae are sister-group to Myodocarpaceae.

Mackinlayoideae G. M. Plunkett et Lowry in South Afr. J. Bot 70(3): 379. 2004

10/c 105. Actinotus (18; Australia, New Zealand), Apiopetalum (2; New Caledonia), Mackinlaya (5; East Malesia, western Pacific islands), Chlaenosciadium (1; C. gardneri; Western Australia), Brachyscias (1; B. verecundus; Western Australia), Xanthosia (c 25; Australia, with their highest diversity in southwestern Western Australia), Pentapeltis (1; P. peltigera; southwestern Western Australia), Schoenolaena (2; Western Australia), Micropleura (2; Colombia, Chile), Centella (c 50; southern Africa to Zimbabwe and Malawi, one species, C. asiatica, pantropical). – Africa, East Malesia, Australia, western South America, with their highest diversity in Western Australia, Melanesia and South Africa. Usually trees or shrubs (rarely herbs). Phellogen in Centella subepidermal. Vessel elements often with scalariform perforation plates. Axial parenchyma usually apotracheal (sometimes also paratracheal). Leaves in Mackinlaya irregularly palmately compound. Inflorescence paniculate or raceme-like, often consisting of compound umbels. Pedicel usually articulated. Sepals in Pentapeltis and Schoenolaena petaloid with ribs containing indistinct oil ducts; sepals absent in Centella and Micropleura. Petals with valvate aestivation, distinctly clawed. Nectary usually divided (in Actinotus present on style). Pistil composed of two (to four) carpels. Fruit usually laterally compressed (not so in Apiopetalum). Wings (“pseudo-wings”), if present, formed by compression or folding of carpel and composed of both mesocarp and endocarp; vascular bundle present at margin. Endocarp lignified. Vittae absent. Carpophore present in, e.g., Mackinlaya; absent in Actinotus and Apiopetalum and in Centella and Xanthosia clades (these having single or paired ventral bundles fused with mericarps). Ribs with distinct oil ducts in, e.g., Actinotus, Chlaenosciadium, Centella, Micropleura, and Xanthosia (in Centella distinct or indistinct). Fruit in Actinotus monocarpellate with lignified endocarp, but without vittae. Centellose (oligosaccharide) present in Centella. x = 10.

[Platysace+[[Klotzschia+[Diposis+Azorelloideae]]+[Hermas+[Saniculoideae+[[Lichtensteinia+[Choritaenia+Marlothiella]]+Apioideae]]]]]

Vessel elements usually with simple perforation plates. Umbels usually compound. Fruit usually dorsally compressed. Carpophore present. Schizocarp with mericarps separating at maturity.

Platysace clade

1/c 25. Platysace (c 25; Australia, with their highest diversity in southwestern Western Australia). – Perennial herbs and suffrutices. Ribs sometimes with oil ducts. Sepals absent. Cotyledons rounded, serrate. n = 8.

[[Klotzschia+[Diposis+Azorelloideae]]+[Hermas+[Saniculoideae+[[Lichtensteinia+[Choritaenia+Marlothiella]]+Apioideae]]]]

Mesocarp vittae irregular, anastomosing and/or branching.

[Klotzschia+[Diposis+Azorelloideae]]

Klotzschia clade

1/3. Klotzschia (3; Brazil). – Pollen grains resembling those in Araliaceae: small size, long colpus, short os, thin exine, similar sexine stratification, and weak differentiation into tectum and short columellae. Fruit without wings or “pseudo-wings”. Ribs without distinct oil ducts. Single fused ventral bundle replacing carpophore.

[Diposis+Azorelloideae]

Diposis clade

1/3. Diposis (3; southern Chile and Argentina). – Carpophore free. Ribs without distinct oil ducts.

Azorelloideae G. M. Plunkett et Lowry in South Afr. J. Bot. 70(3): 379. 2004

15–18/c 95. Homalocarpus (6; Chile), Bolax (4–5; temperate regions in Chile and Argentina), Dichosciadium (1; D. ranunculaceum; Australia), Bowlesia (c 15; South America; incl. Drusa?), Drusa (1; D. glandulosa; the Canary Islands, Somalia; in Bowlesia?), Domeykoa (4; Peru, northern Chile), Eremocharis (9; Peru, Chile), Oschatzia (2; southeastern Australia), Pozoa (2; the Andes in Chile and Argentina), Asteriscium (8; Chile, Argentina), Gymnophyton (6; the Andes in Chile and Argentina), Spananthe (1; S. paniculata; the Andes), Dickinsia (1; D. hydrocotyloides; southwestern China), Diplaspis (3; southeastern Australia, Tasmania), ‘Huanaca’ (4; southern Chile and Argentina; paraphyletic; incl. Schizeilema and Stilbocarpa?), Schizeilema (1; S. frageosum; southeastern Australia; in Huanaca?), Stilbocarpa (1; S. polaris; New Zealand, Stewart Island and adjacent islands, the Antipode Islands, Campbell Island, Auckland Islands, Macquarie Island; in Huanaca?), ’Azorella’ (c 25?; New Zealand, Macquarie Island, from the northern Andes to southern South America, Falkland Islands, subantarctic islands; non-monophyletic; incl. Mulinum?), ‘Mulinum’ (c 20; southern Andes in Chile and Argentina; polyphyletic; in Azorella?). – Southwestern China, Australia, Tasmania, New Zealand, South America, subAntarctic islands, with their highest diversity in the southern Andes; Drusa glandulosa (Bowlesia glandulosa) in the Canary Islands and Somalia. Phellogen in Mulinum deeply seated. Leaves usually simple (rarely compound). Stipules present. Megasporocytes two to four. Megasporangium large and relatively persistent. Megagametophyte tetrasporous, 16-nucleate, Drusa type (and other developmental types). Fruit with wings (“pseudo-wings”) and/or ribs. Wings (“pseudo-wings”), if present, formed by compression or folding of carpel and composed of both mesocarp and endocarp; vascular bundle present at margin. Carpophore present in Schizeilema ranunculus, Stilbocarpa and Huanaca. Carpophores free in Drusa, Homalocarpus, Asteriscium, Gymnophyton, Spananthe, Dickinsia, and Diplaspis (several other genera have fused ventral bundles). Outer mesocarp in Azorella with druses. Companion cells (oil canals associated with vascular bundles) present. Inner layer of endocarp fibres longitudinally arranged. n = 8–10. Petroselinic acid absent at least in Bowlesia. – The clade [Dickinsia+Diplaspis] is sister to the Azorella clade (Nicolas & Plunkett 2012).

[Hermas+[Saniculoideae+[[Lichtensteinia+[Choritaenia+Marlothiella]]+Apioideae]]]

Stipules absent. Ovules incompletely tenuinucellate. Nucellar cap present. Funicle long. Fruit with lateral secondary ribs. Mesocarp non-lignified. Endocarp consisting of single cell layer. Intrajugal vittae (oil ducts in ribs) present.

Hermas clade

1/8. Hermas (8; Western and Eastern Cape). – Perennial herbs or suffrutices. Umbels congested. Carpophore present. Single rhomboidal crystals present in mesocarp cells. Druses absent. Intrajugal vittae present. Vallecular vittae absent. Rib oil ducts present. Endocarp woody. n = 7.

[Saniculoideae+[[Lichtensteinia+[Choritaenia+Marlothiella]]+Apioideae]]

Stipules absent. Basal leaf with pinnate venation. Ovules more or less tenuinucellar. Nucellar cap present. Fruit wings, if present, formed by expansion of mesocarp, consisting only of mesocarp tissue; vascular bundle usually at base. Secondary (lateral) ribs sometimes present. Mesocarp cell walls lignified. Endocarp one cell layer thick, parenchymatous. Cell walls sometimes lignified. Calciumoxalate druses present in mesocarp and around commissure. Rhomboidal crystals absent.

Saniculoideae Burnett Outl. Bot.: 774. Jun 1835

10/330–350. Mainly temperate and subtropical regions; some species in tropical areas. Phellogen sometimes outer cortical. Peripheral stem collenchyma often very well developed. Leaves usually simple (rarely compound). Flowers usually in capitula or simple umbel (inflorescence probably consisting of reduced compound umbels). Nectary sometimes extrastaminal. Style separated from disc by narrow groove. Fruit sessile, usually spiny or scaly. Carpophore not free, or absent. Intrajugal, irregular, vittae (secretory canals/cavities, branched or anastomosing) usually present. Regular vittae absent. Druses usually frequent, present not only in commissural side. Mesocarp vittae present or absent. Ribs with large oil ducts or cavities. Endocarp often lignified. Cotyledons round. n = 8 (9, 11, 12). Kaurene type terpenoids, 3’-O-β-D-glucopranosyl rosmarinic acid (rosmarinic acid glucoside), and cardenolides (in Eryngium) present.

Saniculeae W. D. J. Koch in Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12(1): 66, 138. 1824

7/325–350. Alepidea (c 40; eastern tropical and southern Africa), Actinolema (2; eastern Mediterranean), Arctopus (3; Northern, Western and Eastern Cape), Astrantia (8–9; Central and South Europe, western Asia), Eryngium (c 230–250; temperate to tropical regions on both hemispheres; paraphyletic?), Petagnaea (1; P. gussonii; Sicily), Sanicula (c 40; almost cosmopolitan, absent from New Guinea and Australia). – Cosmopolitan. Usually perennial herbs (rararely shrubs or small trees). Leaf margin often spinose-dentate (with hairy or spiny tooth apices). Inflorescence often with large (involucral) bracts; pseudanthium often present. Flowers (at least female flowers) usually sessile. Fruit usually scaly or spiny (in Petagnaea glabrous). Free carpophore absent. Regular vittae rarely present (few species in Alepidea). Rib oil ducts sometimes absent. Druses usually scattered throughout mesocarp. Endocarp not lignified. n = 8 (9, 11, 12). – Eryngium may be divided in one New World and one Old World clade. Some species of Eryngium have fruit wings entirely consisting of exocarp.

[Phlyctidocarpa+Steganotaenieae]

Phlyctidocarpeae Magee, C. I. Calviño, M. Liu, S. R. Downie, P. M. Tilney et B.-E. van Wyk in Taxon 59: 578. 4 Apr 2010

1/1. Phlyctidocarpa (1; P. flava; Outjo and Kaokoveld in Namibia). – Herbs. Umbels pedunculate, compound. Fruit with bristles and surface vesicles. Ribs bifurcate in cross-section. Regular vittae present. Rib oil ducts large. Druses scattered throughout mesocarp. Ventral bundles weakly developed. n = ?

Steganotaenieae C. I. Calviño et S. R. Downie in Mol. Phylogen. Evol. 44(1): 187. Jan 2007

2/4. Steganotaenia (3; Ethiopia to southern Africa), Polemanniopsis (1; P. marlothii; Northern and Western Cape). – Ethiopia to southern Africa. Shrubs. Phelloderm with chambered crystalliferous cells; phelloderm in Steganotaenia with axial secretory canals. Dilatation of secondary phloem by expansion of axial parenchyma. Leaves proteranthous (developing prior to floral shoots). Fruit heteromericarpous, with two or three exocarp and mesocarp wings. Rib oil ducts forming cavities. Secretory (oil) cavities (intrajugal cavities) associated with ribs strongly expanded. Carpophore present. Mesocarp usually with calciumoxalate druses. Endocarp slightly lignified. n = ?

[[Lichtensteinia+[Choritaenia+Marlothiella]]+Apioideae]

[Lichtensteinia+[Choritaenia+Marlothiella]]

Ribs with oil ducts or cavities.

Lichtensteinieae Magee, C. I. Calviño, M. Liu, S. R. Downie, P. M. Tilney et B.-E. van Wyk in Taxon 59: 578. 4 Apr 2010

1/8. Lichtensteinia (8; Western and Eastern Cape, KwaZulu-Natal, one species on St. Helena). – Annual or perennial herbs, sometimes woody at base. Leaves deciduous, often proteranthous (developing prior to floral shoots). Inflorescence compound umbel. Fruit usually heteromericarpous. Calciumoxalate crystals present in pericarp. Rib oil ducts very large. Irregular vittae present below veins in ribs. Intrajugal (vallecular) vittae absent. Druses scattered throughout mesocarp.

[Choritaenia+Marlothiella]

Choritaenieae Magee, C. I. Calviño, M. Liu, S. R. Downie, P. M. Tilney et B.-E. van Wyk in Taxon 59: 578. 4 Apr 2010

1/1. Choritaenia (1; C. capensis; southern Africa). – Annual herb. Inflorescence compound umbel. Fruit hairy, dorsally compressed, with “inter-rib” wings; marginal wings composed entirely of mesocarp. Mericarps not separating. Mesocarp lignified. Endocarp woody, with globular oil vesicles in wings (instead of normal vittae or rib oil ducts). Druses absent. Carpophore very short, hygroscopic.

Marlothielleae Magee, C. I. Calviño, M. Liu, S. R. Downie, P. M. Tilney et B.-E. van Wyk in Taxon 59: 578. 4 Apr 2010

1/1. Marlothiella (1; M. gummifera; coastal areas in Namibia). – Small shrub. Inflorescence compound umbel. Fruit heteromericarpous, with unicellular stellate hairs. Wings absent. Rib oil ducts very large. Regular vittae absent. Druses scattered throughout mesocarp. Ventral vascular bundles poorly developed or absent.

Apioideae Seem., Fl. Vit.: 112. Jan 1866 [‘Apiaceae’] (mainly according to Downie & al. 2010)

410–420/2.880–2.980. Cosmopolitan, although mainly in northerly temperate regions. Secretory canals/cavities distinct. Leaves usually compound. Stipules usually absent (Foeniculum and some other genera with stipules). Inflorescence usually compound umbel. Outer flowers of umbel often zygomorphic. Stylopodium without groove. Ovule tenuinucellar. Hypostase present. Carpophore free, bifid, with mericarps attached to apex. Regular vittae present, often distinctly vallecular and commissural, or cyclic. Intrajugal oil ducts small or absent. Endocarp one cell layer thick, unlignified. Druses sometimes only on commissural side, or absent. x = 11. Flavones, methylated flavonoids, furanocoumarins, and phenylpropenes present. – The phylogeny and subdivision below follows Downie & al. (2010).

Annesorhizeae Magee, C. I. Calviño, M. Liu, S. R. Downie, P. M. Tilney et B.-E. van Wyk in Taxon 59: 578. 4 Apr 2010

c 6/c 20. ’Annesorhiza’ (c 12; southern Africa; non-monophyletic), Astydamia (1; A. latifolia; the Canary Islands, Madeira), ’Chamarea’ (≥5; western, central and southern South Africa eastwards to Eastern Cape; non-monophyletic), Ezosciadium (1; E. capense; Eastern Cape), Itasina (1; I. filifolia; Western Cape, southwestern Eastern Cape), Molopospermum (1; M. peloponnesiacum; the Pyrenees, the Cevennes, the Alps). – Southern and Central Europe (Molopospermum), Macaronesia (Astydamia), Africa, with their largest diversity in the Cape Provinces. Usually perennial (in Ezosciadium annual) herbs (sometimes lignified). Leaves compound, proteranthous (developing prior to floral shoots) or deciduous. Vascular bundles of fruit strongly lignified. Fruit in Annesorhiza and Molopospermum heteromericarpous. Druses usually scattered throughout mesocarp (in Ezosciadium restricted to commissure). n = 11 (Annesorhiza), 12 (Itasina).

[Heteromorpheae+the remaining Apioideae]

Tanniniferous epidermal cells absent. Druses present only on commissural side of mericarp, or absent.

Heteromorpheae M. F. Watson et S. R. Downie in Amer. J. Bot. 87(2): 289. 2000

11/35. Heteromorpha clade Anginon (12; southern Namibia, Northern, Western and Eastern Cape), Heteromorpha (7; Africa south of Sahara, Socotra, Yemen), Polemannia (3; mountains in South Africa and Lesotho). – Malagasy clade Andriana (3; Madagascar), Anisopoda (1; A. bupleuroides; Madagascar), Cannaboides (2; Madagascar), Dracosciadium (2; KwaZulu-Natal), Oreofraga (1; O. morrisiana; Socotra), Pseudocannaboides (1; P. andingitrensis; Madagascar), Pseudocarum (2; tropical East Africa, Madagascar), Tana (1; T. bojerianum; Madagascar). – Africa, Madagascar, Socotra, Yemen, with their largest diversity in southwestern Africa. Trees, shrubs, suffrutices or lianas (Dracosciadium herbs with persistent leaves). Walls of vessel elements with helical thickenings. Fibres septate. Sepals sometimes well developed. Fruit not or only slightly dorsiventrally or laterally compressed, in Heteromorpha heteromericarpous. Vittae in Pseudocarum branching and anastomosing. Druses scattered throughout mesocarp.

[Bupleureae+the crown Apioideae]

Druses absent.

Bupleureae Spreng. in J. J. Roemer et J. A. Schultes, Syst. Veg. 6: xxxiii. Aug-Dec 1820 [‘Bupleurinae’]

1/180–190. Bupleurum (180–190; Europe, the Canary Islands, the Mediterranean, northern Africa, temperate Asia, one species, B. mundii, in South Africa, one species, B. americanum, in arctic North America). – Usually annual or perennial herbs (rarely shrubs with secondary lateral growth). Walls of vessel elements with helical thickenings. Fibres septate. Leaves simple, often grass-like. Leaf margin entire. Venation often parallel. Pollen grains usually rhomboidal. Vittae branching and anastomosing. Cotyledons linear, with single vein, glabrous. n = 7, 8.

The “Crown Apioideae

Pericarp without druses.

[Pleurospermopsis clade+the rest]

Pleurospermopsis clade

1/1. Pleurospermopsis (1; P. sikkimense; Bhutan, Nepal, Sikkim, eastern Himalaya, western China). – Biennial or perennial herb. Leaves pinnately compound. Inflorescence compound umbel. Bracts (involucral bracts) and bracteoles (involucel bracts) prominent. Fruit laterally slightly compressed. Ribs prominent, narrowly winged. Carpophore bipartite.

[Chamaesium clade+the rest]

Chamaesium clade

1/5. Chamaesium (5; Central Asia, Himalayas, northern India, Tibet, southwestern China). – Perennial herbs. Leaf pinnately compound. Bracts and bracteoles few or absent. Fruit smooth. Ribs prominent to narrowly winged. Carpophore divided to base.

[Diplolophium clade+the rest]

Diplolophium clade

1/5–7. Diplolophium (5–7; tropical to southern Africa). – Perennial herbs. Leaves pinnately compound. Bracts and bracteoles present. Carpophore divided to base.

[Pleurospermeae+the rest]

Pleurospermeae M. F. Watson et S. R. Downie in Amer. J. Bot. 87: 289. 15 Feb 2000

c 10/68. Aulacospermum (14; Europe, temperate Asia), Pleurospermum (2; Europe, temperate Asia), Trachydium (1; T. roylei; Himalayas), Pseudotrachydium (5; southwestern Asia), Hymenidium (c 35; Pakistan and eastwards to China), Eleutherospermum (1; E. cicutarium; Caucasus, southwestern Asia), Physospermum (2; Europe, temperate Asia), Eremodaucus (1; E. lehmannii; Caucasus to Central Asia and Afghanistan), Korshinskya (5; southwestern and Central Asia), ‘Physospermopsismuliensis (southwestern Sichuan, northwestern Yunnan), ‘Physospermopsisshaniana (Sichuan, Xizang, Yunnan, Burma). – Temperate and alpine regions in Eurasia.

[[Physospermopsis clade+Komarovia clade]+rest]

Physospermopsisclade (East Asia clade)

c 15/57–58. Hansenia (3; northern and Central Asia), Haplosphaera (2; southern and eastern Asia), Heptaptera (6; eastern Mediterranean, southwestern Asia), Hymenolaena (3; Central Asia, Himalayas), Keraymonia (4; Himalayas),‘Notopterygium’ (6; China; non-monophyletic), ‘Physospermopsis’ (14–15; Central to East Asia; non-monophyletic), ‘Pimpinella’ pro parte (3),‘Sinocarumcruciatum (western China, northern Burma), ‘Sinocarumdolichopodum (western Sichuan, northwestern Yunnan), Sinolimprichtia (1; S. alpina; eastern Tibet), ‘Spuriopimpinellabrachycarpa (southeastern Russia, China, northern Korean Peninsula), ‘Tongoloa’ (10–15; Central Asia to northern India; non-monophyletic), ‘Trachydiumsimplicifolium (northwestern Yunnan), ‘Vicatiabipinnata (northwestern Sichuan, Yunnan). – Temperate and alpine regions in Asia.

Komarovieae J. Zhou et S. R. Downie in Mol. Phylogenet. Evol. 53(1): ?. 6 Jun 2009

7/10. Parasilaus (2; southwestern and Central Asia), Komarovia (1; K. anisoptera; Central Asia), Calyptrosciadium (1; C. polycladum; Iran, Afghanistan), Changium (2; Tibet, eastern China), Chuanminshen (1; C. violaceus; China), Cyclorhiza (2; southwestern China), Sphaerosciadium (1; S. denaense; Central Asia). – Southwestern Asia to China.

[Erigenieae+the rest]

Erigenieae Rydb. ex Pimenov et Constance in Taxon 34: 497. 19 Aug 1985

1/1. Erigenia (1; E. bulbosa; eastern United States). – Small glabrous perennial herb. Corm present. Vesicular-arbuscular mycorrhiza absent. Leaves ternately compound. Involucral bracts absent. Bracteoles (involucel bracts) present. Fruit slightly compressed.

[Oenantheae+the rest]

Oenantheae Dumort., Fl. Belg.: 79. 1827

c 20/c 175. ’Sium’ (9; the Northern Hemisphere, Africa; polyphyletic), Berula (7; temperate regions on the Northern Hemisphere, eastern and southern Africa, St. Helena), Cryptotaenia (6; temperate regions on the Northern Hemisphere, East African mountains), Helosciadium (c 45; temperate regions in the Old World), Cicuta (8; temperate regions on the Northern Hemisphere), Oenanthe (c 40; temperate regions on the Northern Hemisphere, tropical African mountains, India to Malesia and Australia), Oxypolis (7; North America), Neogoezia (5; Mexico), Atrema (1; A. americanum; southern United States), Trepocarpus (1; T. aethusae; southern United States), Daucosma (1; D. laciniatum; the United States), Lilaeopsis (c 20; Mauritius, Australia, New Zealand, North and South America), Cynosciadium (2; southern United States), Limnosciadium (2; southern central United States), Harperella (1; H. nodosa; estern United States), Tiedemannia (2; eastern United States, Cuba), Ptilimnium (4–5; eastern United States), Perideridia (13; North America), Trocdaris (1; T. verticillata; western Europe, Morocco). – Cosmopolitan.

[apioid superclade+Aciphylleae+Acronema clade+Arcuatopterus clade+Conioselinum chinense clade+Scandiceae+Smyrnieae]

Aciphylleae M. F. Watson et S. R. Downie in Amer. J. Bot. 87: 288. 15 Feb 2000

c 5/c 70. ‘Aciphylla’ (c 40; eastern Australia, Tasmania, New Zealand; non-monophyletic; incl. Anisotome?),‘Anisotome’ (16; New Zealand, subAntarctic islands; non-monophyletic; in Aciphylla?), Lignocarpa (2; New Zealand; in Aciphylla?), Scandia (2; New Zealand; in Aciphylla?), Gingidia (10; Australia, Tasmania, New Zealand; in Aciphylla?). – Mainly New Zealand, some species of ‘Aciphylla’ in eastern Australia.

Acronema clade

15–20/150–160. Acronema (c 25; Himalayas, western China), ’Angelicaanomala (Siberia, China, the Korean Peninsula), Halosciastrum (1; H. melanotilingia; East Asia), Harrysmithia (2; China), Kitagawia (5; Central Asia), ‘Ligusticum’ (40–50; temperate regions on the Northern Hemisphere; non-monophyletic), Meeboldia (5; Himalayas), Oreocomopsis (3; Himalayas, southern China), Ostericum grosseserratum (Mongolia, China, the Korean Peninsula), Ostericum scaberulum (Yunnan), Pachypleurum (2; temperate regions in Europe and Asia), ‘Pleurospermumhookeri (China, Nepal, Sikkim, Bhutan), ‘Pleurospermumyunnanense (western Sichuan, northwestern Yunnan, northeastern Burma), Pternopetalum (15; eastern Nepal, Bhutan, China, southern Japan), Pterygopleurum (1; P. neurophyllum; the Korean Peninsula, southern Japan), Rupiphila (1; R. tachiroei; China), Sinocarum (c 25; China), Spuriopimpinella (3; China), ‘Tilingia’ (3; China, northeastern Asia, Alaska; non-monophyletic). – Temperate and alpine regions in Eurasia, especially Central Asia to China.

Arcuatopterus clade

1/6. Arcuatopterus (6; eastern Himalaya, southwestern China). – Glabrous perennial herbs. Leaves two- or three-pinnatisect. Bracts usually absent (sometimes one, caducous). Bracteoles absent. Fruit dorsally strongly compressed. Dorsal ribs indistinct. Lateral ribs broadly winged; wings thin to corky. Carpophore divided.

Conioselinum chinense clade

c 5/11. ‘Conioselinumchinense (Russia, China, Japan, North America), ‘Conioselinumscopulorum (western United States), ‘Ligusticumcanadense (North America), ‘Ligusticumporteri (western United States, northern Mexico), Meum (3; Europe, northern Africa), Mutellina (3; temperate regions in Europe and Asia), Trochiscanthes (1; T. nodiflora; southern Europe). – Temperate regions on the Northern Hemisphere.

Scandiceae Spreng. in J. J. Roemer et J. A. Schultes, Syst. Veg. 6: xliii. Aug-Dec 1820 [‘Candicinae’]

c 40/c 335. Scandicinae Tausch in Flora 17: 342. 14 Jun 1834. Conopodium (6; Europe, the Mediterranean, temperate regions in Asia), Athamanta (7; Europe, the Canary Islands, the Mediterranean), Todaroa (1; T. aurea; the Canary Islands), Sphallerocarpus (1; S. gracilis; southern Asia), ’Chaerophyllum’ (33; temperate regions on the Northern Hemisphere; paraphyletic), Scandix (7; Europe, the Mediterranean), Osmorhiza (10; East Asia, America), Myrrhis (1; M. odorata; Europe), Anthriscus (9; Europe, mountains in Africa, temperate Asia), Geocaryum (13–15; eastern Mediterranean), Kozlovia (4; Central Asia to India), Chaerophyllopsis (1; C. huai; western China). – Ferulinae Drude in H. G. A. Engler et K. A. E. Prantl, Nat. Pflanzenfam. III, 8: 115. Dec 1897. Ferula (c 130; the Canary Islands, the Mediterranean and eastwards to Central Asia), Autumnalia (2; southwestern and Central Asia)?, Fergania (1; F. polyantha; Central Asia)?, Kafirnigania (1; K. hissarica; Central Asia)? – Torilidinae Dumort., Fl. Belg.: 81. 1827. Astrodaucus (2; Europe, temperate regions in Asia), Glochidotheca (1; G. foeniculacea; Europe, western Asia), Szovitsia (1; S. callicarpa; Caucasus, Armenia, Iran), Torilis (15; Europe, the Canary Islands, the Mediterranean and eastwards to East Asia, tropical and southern Africa), Yabea (1; Y. microcarpa; western United States, northwestern Mexico), Caucalis (1; C. platycarpos; southern Europe), Turgenia (1; T. latifolia; Central Europe, the Mediterranean and eastwards to Central Asia), Lisaea (3; eastern Mediterranean, southwestern Asia). – Daucinae Dumort., Fl. Belg.: 81. 1827. ‘Laserpitium’ (c 35; Europe, the Canary Islands, the Mediterranean, southwestern Asia; non-monophyletic), Polylophium (2; western Asia), Thapsia (14; the Iberian Peninsula, the Canary Islands, the Mediterranean, northern to tropical Africa), Laser (1; L. trilobum; central and southern Europe, western Asia), Cuminum (4; the Mediterranean to Sudan and Central Asia), Orlaya (3; southeastern Europe to Central Asia), Monizia (1; M. edulis; Madeira, the Salvage Islands), Daucus (22; Europe, the Mediterranean, North and tropical Africa, southwestern and Central Asia, Australia, New Zealand, America), ‘Athamantadella-cellae (Libya; in Daucus?) ‘Cryptotaeniaelegans (the Canary Islands), Melanoselinum (1; M. decipiens; Madeira), Tornabenea (4; the Cape Verde Islands). – Artedia cladeArtedia (1; A. squamata; eastern Mediterranean). – Glaucosciadium clade Glaucosciadium (1; G. cordifolium; southern Turkey, Cyprus). – Scandicinae is distributed mainly in warm-temperate regions on the Northern Hemisphere; the Ferula clade in southwestern to Central Asia, with few species in the Mediterranean and Macaronesia; Torilidinae in temperate and warmer regions in the Old World, with one species in western North America; Daucinae mainly in warm-temperate and subtropical regions on the Northern Hemisphere, with their highest diversity in Macaronesia, the Mediterranean and southwestern Asia; and the Artedia and the Glaucosciadium clades in the eastern Mediterranean.

Smyrnieae Spreng. in J. J. Roemer et J. A. Schultes, Syst. Veg. 6: xxxvii. Aug-Dec 1820 [’Smyrniae’].

2/8. Smyrnium (7; southern and southeastern Europe, the Mediterranean), Lecokia (1; L. cretica; Crete and eastwards to Iran). – Southern and southeastern Europe, the Mediterranean, southwestern Asia.

The Apioid Superclade

Apieae Takht. ex V. M. Vinogr., Fl. Vostochnoĭ Evropy 11: 339. 2004

12–13/34–38. Apium (3–5?; cosmopolitan), Naufraga (1; N. balearica; Majorca in Apium?), Anethum (1; A. graveolens; southwestern Asia?), Ridolfia (1; R. segetum; the Mediterranean), Pseudoridolfia (1; P. fennanei; Morocco), Foeniculum (4–5; temperate regions in Asia), Deverra (10; arid and semi-arid regions in North Africa to southern Africa, the Arabian Peninsula and southwestern Asia), Ammi (3–4; Macaronesia, the Mediterranean, northern Africa, western Asia), Petroselinum (2; Europe, the Mediterranean), Billburttia (2; Madagascar), Sclerosciadium (1; S. nodiflorum; Europe, the Mediterranean, Africa), ‘Seseliwebbii (the Canary Islands), Stoibrax (4; the Iberian Peninsula, North Africa). – Cosmopolitan, with their highest diversity in warm-temperate regions in Africa to southwestern Asia. – Naufraga balearica lacks free carpophore and probably has lignified endocarp.

Cachrydinae Meisn., Plant. Vasc. Gen.: Tab. Diagn. 150, Comm. 108. 16-22 Sep 1838 [’Cachrydeae’]

8/100–105. Alococarpum (1; A. erianthum; Iran), Azilia (1; A. eryngioides; Iran), Bilacunaria (4; southwestern Asia), Cachrys (3–4; Europe, North Africa), Diplotaenia (2; Turkey, Iran), Ekimia (1; E. bornmuelleri; Turkey), Eriocycla (7; northern Iran, western Himalaya to northern and western China), Ferulago (c 45; southeastern Europe, the Mediterranean and eastwards to Central Asia), Prangos (38; eastern Europe, eastern Mediterranean, temperate Asia). – Warm-temperate regions in Europe, North Africa and Asia, with their largest diversity in the eastern Mediterranean to southwestern Asia. – Bilacunaria has oil vesicles dispersed in mesocarp.

Careae Baill., Hist. Plant. 7: 174, 219. Apr 1879

10–11/37–38. Carum (5; Europe, Caucasus, southwestern Asia), Fuernrohria (1; F. setifolia; Caucasus, Armenia; in Carum?), ‘Grammosciadium’ (7; eastern Mediterranean; non-monophyletic; in Carum?), Chamaesciadium (1; C. acaule; western Asia; in Carum?), Rhabdosciadium (5; eastern Europe, western Asia), Falcaria (4–5; Central Europe, the Mediterranean, western and Central Asia), Aegokeras (1; A. caespitosa; Turkey), ‘Aegopodium’ (10; temperate regions in Europe and Asia; non-monophyletic), Gongylosciadium (1; G. falcarioides; Turkey, Caucasus, Iran), Hladnikia (1; H. pastinacifolia; Slovenia), Pseudopimpinella (1; P. anthriscoides; Iran). – Temperate regions in Eurasia, with the higest diversity in southwestern to Central Asia.

Coninae Rouy, Consp. Fl. France: 116. 15 Aug 1927 [‘Conieae’]

1/6. Conium (6; southern Europe, the Mediterranean, temperate Asia, southern Africa).

Coriandreae W. D. J. Koch in Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12(1): 65, 82. 1824

2/6. Coriandrum (3; southeastern Europe, southwestern Asia), Bifora (3; the Mediterranean and eastwards to Central Asia, southern United States, northern Mexico?). – Southeastern Europe, the Mediterranean, southwestern to Central Asia, one species of Bifora (B. americana) in the southern United States and northern Mexico?

Echinophoreae Benth. et Hook. f., Gen. Plant. 1: 862, 865. Sep 1867

c 9/31. Rughidia (2; Yemen), Dicyclophora (1; D. persica; Iran), Pycnocycla (12; tropical West Africa and eastwards to northwestern India), ‘Echinophora’ (9; the Mediterranean, southwestern Asia to Iran; non-monophyletic), Anisosciadium (3; southwestern Asia), Mediasia (1; M. macrophylla; southwestern and Central Asia), Nirarathamnos (1; N. asarifolius; Haggeher Mountains on Socotra), ’Pimpinellaheyneana (Sri Lanka), ’Trachyspermumaethusifolium (Somalia). – Warm-temperate to subtropical regions in West Africa and the Mediterranean to Central and South Asia.

Opopanax clade

7/11. Smyrniopsis (1; S. aucheri; eastern Mediterranean, southwestern Asia to Iran), Opopanax (3; the Balkans and eastwards to Iran), Crenosciadium (1; C. siifolium; Turkey), Krubera (1; K. peregrina; western Mediterranean), Magydaris (2; the Mediterranean), Petroedmondia (1; P. syriaca; southwestern Asia), Stefanoffia (2; eastern Mediterranean). – The Mediterranean to southwestern Asia. – Smyrniopsis has oil vesicles dispersed in mesocarp. Krubera is sister to Coriandreae (with Levisticum sister to these three genera) in the analyses by Magee & al. (2009). The exact placement of Krubera is very uncertain at this moment and has to await phylogenetic investigations including a wide spectrum of taxa.

Pimpinelleae Spreng. in J. J. Roemer et J. A. Schultes, Syst. Veg. 6: xxxiv. Aug-Dec 1820

15–17/c 235. ’Psammogeton’ (7; southwestern Asia; non-monophyletic), ’Aphanopleura’ (6; Central Asia to Afghanistan; non-monophyletic), ’Pimpinella’ (c 200; Europe, Macaronesia, the Mediterranean, temperate Asia, East African mountains; non-monophyletic), Arafoe (1; A. aromatica; Caucasus), Bubon (1; B. macedonicum; southern Italy, southern Balkan), ‘Cryptotaeniaafricana (Central and East Africa), Demavendia (1; D. pastinacifolia; southwestern and Central Asia), Frommia (1; F. ceratophylloides; southern Central Africa), Haussknechtia (1; H. elymaitica; southwestern Iran), Nothosmyrnium (2; East Asia), Opsicarpium (1; O. insignis; Iran), Phellolophium (3; Madagascar), ’Physospermopsiscuneata (Sichuan, Yunnan), ’Seselidiffusum (northern India), ’Trachyspermumscaberulum (western China), ’Trachyspermumtriradiatum (Sichuan), ’Zeravschania’ (11; southwestern to Central Asia; non-monophyletic). – Temperate to tropical regions in Eurasia and Africa.

Pyramidoptereae Boissier, Fl. Orient. 2: 1089. 1872 [‘Pyramidopterae’]

c 30/175–180. Ammoides (2; the Mediterranean), Astomaea (2; eastern Mediterranean and eastwards to Central Asia), ‘Bunium’ (45–50; Europe, the Mediterranean, North Africa, southwestern and Central Asia; non-monophyletic), ‘Carum’ (c 25?, e.g. C. heldreichii, C. rupestre and C. rupicola; C. buriaticum; C. appuanum and C. graecum; temperate regions on the Northern Hemisphere; polyphyletic), Crithmum (1; C. maritimum; Atlantic coasts of Europe, the Mediterranean), Cyclospermum (1; C. leptophyllum; Central America, the West Indies), Elaeosticta (c 25; eastern Europe, western and Central Asia), Galagania (5; eastern Europe to Central Asia), Gongylotaxis (1; G. rechingeri; Afghanistan), Hellenocarum (3; southeastern Europe, southwestern Asia), ‘Hyalolaena’ (10; southwestern and Central Asia; non-monophyletic), Indoschulzia (2; Himalayas), Kosopoljanskia (3; Central Asia), Lagoecia (1; L. cuminoides; the Mediterranean), Lipskya (1; L. insignis; Central Asia), Mogoltavia (2; Central Asia), Muretia (5; southern Russia, western and Central Asia), Notiosciadium (1; N. pampicola; Argentina),‘Oedibasis’ (4; Central Asia; non-monophyletic), Oreoschimperella (3; mountains in Ethiopia, Kenya and Yemen), Ormopterum (2; Central Asia, Pakistan), ‘Pimpinellasiifolia (western Pyrenees, northern Spain), Postiella (1; P. capillifolia; Turkey), Pyramidoptera (1; P. cabulica; Afghanistan), Scaligeria (3; the Balkan, southwestern Asia), Schrenkia (12; Central Asia), Schtschurowskia (2; Central Asia), Schulzia (4; Central Asia, northwestern India), Sison (2; western and southern Europe, the Mediterranean), Tamamschjanella (3; the Balkans, Caucasus, southwestern Asia), ‘Trachyspermumammi (northeastern Africa). – Temperate to subtropical regions on the Northern Hemisphere and one species in Argentina, with the largest diversity in the Mediterranean to Central Asia. – Lagoecia has well developed sepals and single-seeded fruit. Notiosciadium pampicola has compound umbels, laterally compressed fruits and entire carpophores.

Selineae Spreng. in J. J. Roemer et J. A. Schultes, Syst. Veg. 6: xlvi. Aug-Dec 1820

c 65/c 730–785. Aethusa (1; A. cynapium; Europe, North Africa, western Asia), Ammoselinum (3; North America, temperate South America), ‘Angelica’ (c 110; temperate regions on the Northern Hemisphere; non-monophyletic), Apiastrum (2; North America), ‘Cervaria’ (4; temperate regions in Europe and Asia; non-monophyletic), ‘Cortia’ (8; Central Asia, Himalayas; non-monophyletic), Cortiella (3; Central and South Asia), Dichoropetalum (c 30; southern and southeastern Europe, the Mediterranean, North Africa, southwestern Asia), Dimorphosciadium (1; D. gayoides; Central Asia), Dystaenia (2; the Korean Peninsula, Japan), Endressia (2; the Pyrenees, northern Spain), Exoacantha (1; E. heterophylla; southwestern Asia), Ferulopsis (2; northern and Central Asia), Glehnia (2; northeastern Asia, western North America), Imperatoria (3; Europe, North Africa), Johrenia (12; Greece, Turkey, southwestern and Central Asia), Kailashia (2; northern India, Tibet), Karatavia (1; K. kultiassovii; Central Asia), Kedarnatha (5; Himalayas, northern Burma), Ledebouriella (2; Central Asia), Ligusticopsis (14–15; temperate East Asia), ‘Ligusticum’ pro parte (7; temperate regions), Lomatocarpa (4; southwestern and Central Asia, Afghanistan), Lomatocarum (1–2; L. alpinum; Caucasus), Magadania (2; northeastern Asia), Oligocladus (2; Argentina), Oreocome (6; Pakistan to western China), Ormosolenia (1; O. alpina; eastern Mediterranean), ‘Pachypleurummutellinoides (Central Europe, Arctic Russia, northern Ural), Paraligusticum (1; P. discolor; Altai Mountains), ’Peucedanum’ (100–120; temperate regions in Europe and Asia, mountains in tropical and southern Africa; polyphyletic), Pilopleura (2; Central Asia), Pteroselinum (11; Europe, temperate Asia), Rumia (1; R. crithmifolia; Crimea), Sajanella (1; S. monstrosa; southern Siberia), ‘Selinum’ (2; Europe and eastwards to Central Asia; non-monophyletic; incl. Cnidium?), Cnidium (4–5?; in Selinum?), ’Seseli’ (100–120; Europe and eastwards to Central Asia, northern tropical Africa; non-monophyletic), Siculosciadium (1; S. nebrodense; Sicily), Spermolepis (5; North America, Argentina, the Hawaii Islands), Stenocoelium (3; Central Asia), Thecocarpus (2; Turkey, Iran), Tommasinia (1; T. verticillaris; Central Europe), Trinia (8–10; the Mediterranean and eastwards to Central Asia), Vicatia (4; southern Siberia, Central Asia, Himalayas to western China), Xanthoselinum (2; southern and southeastern Europe, Caucasus to western Siberia). – Arracacia cladeArracacia’ (c 55; tropical America; polyphyletic), Coaxana (2; Mexico), Coulterophytum (5; Mexico), Dahliaphyllum (1; D. almedae; Mexico), Donnellsmithia (15–20; Mexico, Central America), Enantiophylla (1; E. heydeana; Central America), Mathiasella (1; M. bupleuroides; Mexico), Neonelsonia (2; Mexico, Central and northern South America), Myrrhidendron (5; Central America, Colombia), Ottoa (1; O. oenanthoides; Mexico), Prionosciadium (8; Mexico), Rhodosciadium (14; Mexico), Cotopaxia (2; Colombia, Ecuador)?, Niphogeton (18; northern Andes), Perissocoeleum (4; Colombia). – Perennial endemic North American cladeLomatium’ (c 75; western North America; polyphyletic), ’Cymopterus’ (32; western North America; polyphyletic; partially in Lomatium), Eurytaenia (2; North America), Podistera (4; western North America), Shoshonea (1; S. pulvinata; Wyoming). – Mainly temperate and alpine regions on both hemispheres. The Arracacia clade is distributed in Mexico, Central and South America. The Perennial endemic North American clade is centered in the Rocky Mountains. – Myrrhidendron, in Central America and Colombia, is woody with secondary lateral growth.

Sinodielsia clade

c 15/c 50. ‘Angelica’ pro parte (3), Cenolophium (1; C. denudatum; eastern Europe, temperate Asia), ‘Cnidiumdahuricum (Siberia), ‘Cnidiumofficinale (China), Conioselinum (18; eastern Europe, temperate regions in Asia and North America), Levisticum (1; L. officinale; eastern Mediterranean), ‘Ligusticumacuminatum (China), ‘Ligusticumjeholense (China), ‘Ligusticumsinense (China), ‘Ligusticumtenuissimum (China, the Korean Peninsula), Lithosciadium (2; Central and northern Asia to western China), Paulita (3; Central Asia), ‘Pleurospermumwrightianum (western China), ‘Prangoshaussknechtii (Iran), Pterocyclus (4; Himalayas, southwestern China), Seselopsis (2; Central Asia), Silaum (1; S. silaus; Europe, temperate Asia), Sinodielsia (2–3; southeastern Tibet, Yunnan), Sphaenolobium (3; Central Asia), ‘Vicatiathibetica (Nepal, eastern and southern Tibet, western China). – Temperate and alpine regions on the Northern Hemisphere, with their highest diversity in Central Asia, the Himalayas and China.

Tordylieae W. D. J. Koch in Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12(1): 65, 85. 1824.

25 – 27/210–215. Tordyliinae Drude in H. G. A. Engler et K. A. E. Prantl, Nat. Pflanzenfam. III, 8: 115. Dec 1897. ‘Angelicalikiangensis (Guizhou, Yunnan), ‘Angelicaoncosepala (northwestern Yunnan), ‘Heracleum’ (c 65; temperate regions on the Northern Hemisphere, East Afican tropical mountains, with their largest diversity in Himalayas and western China; incl. Mandenovia?; non-monophyletic), Mandenovia (1; M. komarovii; Caucasus; in Heracleum?), Kandaharia (1; K. rechingerorum; Afghanistan), Lalldhwojia (2; Himalayas), Malabaila (8–10; the Balkan, Turkey, Caucasus and eastwards to Iran), Leiotulus (9; Europe, temperate Asia), ‘Pastinaca’ (14; Europe, temperate Asia; non-monophyletic), ‘Semenovia’ (18; Asia; non-monophyletic), Tetrataenium (7–8; southwestern to Central Asia), Tordyliopsis (1; T. brunonis; Himalayas, Tibet), Tordylium (18; southern Europe, the Mediterranean, southwestern Asia), Trigonosciadium (4; Turkey, Iraq, Iran), Vanasushava (1; V. pedata; southern India), Zosima (4; southwestern and Central Asia). – Cymbocarpum clade Cymbocarpum (3–4; southwestern Asia). – Lefebvrea clade Afrosciadium (18; eastern and southeastern Africa), Afroligusticum (2; tropical Africa), Capnophyllum (1; coastal areas of Northern and Western Cape), Cynorhiza (3; Western Cape), Dasispermum (7; coastal areas in South Africa), Scaraboides (1; S. manningii; Vanrhynsdorp and Tanqua karoo in Northern and Western Cape), Nanobubon (2; Western Cape, western Eastern Cape), Notobubon (11; Western and Eastern Cape), Stenosemis (2; Eastern Cape to southern KwaZulu-Natal), ‘Notobubonpearsonii (western Northern Cape), Lefebvrea (6; tropical and southwestern Africa). – Tordyliinae are distributed in temperate and alpine regions on the Northern Hemisphere, tropical African mountains and southern India, with their largest diversity in southwestern to Central Asia and the Himalayas; the Cymbocarpum clade occurs in southwestern Asia; and the Lefebvrea clade in tropical to southern Africa, with their highest diversity in the Cape Provinces.

Unplaced Apioideae Adenosciadium (1; A. arabicum; southwestern Arabian Peninsula), Aframmi (2; Angola), Afrosison (3; tropical Africa), Agasyllis (1; A. caucasica; Caucasus), Angoseseli (1; A. mossamedensis; Angola), Apodicarpum (1; A. ikenoi; eastern Japan), Asciadium (1; A. coronopifolium; Cuba), Austropeucedanum (1; A. oreopansii; northwestern Argentina), Bonannia (1; B. graeca; southern Italy, southern Greece), Caropsis (1; C. verticillato-inundata; Europe), Cephalopodum (3; Central Asia), Chabrea (1; C. carvifolia; Europe), Coristospermum (3; Europe), Cyathoselinum (1; C. tomentosum; southeastern Europe), Dethawia (1; D. splendens; the Pyrenees, Cordillera Cantabrica), Ducrosia (5; Egypt to northwestern India), Ergocarpon (1; E. cryptanthum; eastern Iraq, western Iran), Eriosynaphe (1; E. longifolia; southeastern Russia to Central Asia), Erythroselinum (1; E. atropurpureum; Ethiopia), Froriepia (2; Turkey to Iran), Grafia (1; G. golaka; Italy, northwestern Balkan), Haplosciadium (1; H. abyssinicum; Ethiopia to East African mountains), Horstrissea (1; H. dolinicola; Crete), Kalakia (1; K. marginata; Iran), Karnataka (1; K. benthamii; southern India), Kelussia (1; K. odoratissima; Iran), Kundmannia (2; southern Europe, the Mediterranean), Ladyginia (3; southwestern and Central Asia), Macroselinum (1; M. latifolium; southern Europe, Caucasus), Mastigosciadium (1; M. hysteranthum; Afghanistan), Microsciadium (1; M. minutum; Turkey), Oliveria (1; O. decumbens; Syria to Iran), Opoideia (1; O. galbanifera; Iran), Ormosciadium (1; O. aucheri; eastern Turkey), Palimbia (3; southern and eastern Russia to Central Asia), Pancicia (1; P. serbica; western Balkan Peninsula), Paraselinum (1; P. weberbaueri; Peru), Pastinacopsis (1; P. glacialis; Central Asia), Pedinopetalum (1; P. domingense; Hispaniola), Physotrichia (10; tropical Africa), Pinacantha (1; P. porandica; Afghanistan), Pinda (1; P. concanensis; Western Ghats, southern India), Polyzygus (1; P. tuberosus; southern India), Portenschlagiella (1; P. ramosissima; southern Italy, western Balkan), Pseudoselinum (1; P. angolense; Angola), Ptychotis (1; P. saxifraga; Central and southwestern Europe), Registaniella (1; R. hapaxlegomena; Afghanistan), Rhopalosciadium (1; R. stereocalyx; Iran; in Torilis?), Rhysopterus (3; North America), Rouya (1; R. polygama; Corsica, Sardinia, North Africa), Rutheopsis (1; R. herbanica; Lanzarote and Fuerteventura in the Canary Islands), Sciothamnus (4; southern Africa), Sclerochorton (1; S. haussknechtii; Iran), Sclerotiaria (1; S. pentaceros; Central Asia), Scrithacola (1; S. kurramensis; Afghanistan, Pakistan), Selinopsis (2; the Mediterranean), Spuriodaucus (3; tropical Africa), Stenotaenia (5–6; southwestern Asia), Stewartiella (2; Pakistan), Taeniopetalum (2; the Balkan Peninsula, Turkey), Tamamschjania (2; northern Greece, Turkey), Thamnosciadium (1; T. junceum; Greece), Trachyspermum p.p. (8; tropical and northeastern Africa to Central Asia, India and western China), Tricholaser (2; southern and southwestern Asia), Xatardia (1; X. scabra; eastern Pyrenees).

Cladogram (simplified) of Apiaceae based on DNA sequence data (Nicolas & Plunkett 2009).

Cladogram (simplified) of Azorelloideae and some other Apiaceae based on DNA sequence data (Nicolas & Plunkett 2009).

Cladogram of Saniculoideae based on DNA sequence data (Nicolas & Plunkett 2009; Magee & al. 2010).

Phylogeny of Apioideae based on DNA sequence data (Downie & al. 2010).

ARALIACEAE Juss.

( Back to Araliales )

de Jussieu, Gen. Plant.: 217. 4 Aug 1789 [’Araliae’], nom. cons.

Hederaceae Giseke, Prael. Ord. Nat. Plant.: 519. Apr 1792; Hydrocotylaceae Bercht. et J. Presl, Přir. Rostlin: 258. Jan-Apr 1820 [’Hydrocotyleae’], nom. cons.; Hederales Link, Handbuch 2: 5. 4-11 Jul 1829 [’Hederaceae’]; Hydrocotylineae Link, Handbuch 1: 314. 4-11 Jul 1829 [’Hydrocotylinae’]; Botryodendraceae J. Agardh, Theoria Syst. Plant.: 231. Apr-Sep 1858 [’Botryodendreae’]

Genera/species 35–40?/c 2.000

Distribution Mainly tropical and subtropical regions in the Southern and Northern Hemispheres; a few genera in temperate areas.

Fossils Leaves, pollen grains and endocarps of Araliaceae have been reported from Cenozoic strata in Europe and Asia. The oldest known fossils of Araliaceae – endocarps of Acanthopanax and Aralia – have been found in the Maastrichtian of Germany. Fossilized endocarps are also recorded from North America.

Habit Usually bisexual (sometimes monoecious, andromonoecious, gynomonoecious, polygamomonoecious, or dioecious), evergreen or deciduous trees, shrubs or lianas (sometimes perennial herbs or suffrutices). Some species are aquatic or helophytes. Leaf scars large and distinct.

Vegetative anatomy Phellogen ab initio superficial. Peripheral collenchyma? Medullary and cortical vascular bundles often present. Vessel elements with simple and/or scalariform (sometimes reticulate etc.) perforation plates; lateral pits scalariform, opposite or alternate, simple and/or bordered pits. Vestured pits present. Imperforate tracheary xylem elements usually libriform fibres (in Boerlagiodendron fibre tracheids) with simple or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, usually heterocellular (rarely homocellular). Axial parenchyma usually paratracheal scanty, often vasicentric (sometimes apotracheal diffuse), or absent. Tyloses often abundant. Sieve tube plastids S type. Nodes usually ≥5:≥5, multilacunar with five or more leaf traces (sometimes 3:3, trilacunar with three traces). Parenchyma and other tissues with schizogenous secretory canals and cavities containing ethereal oils, resins and gum. Prismatic calciumoxalate crystals sometimes frequent.

Trichomes Hairs unicellular or multicellular, uniseriate or branched, often stellate or dendritic (sometimes T-shaped or peltate-lepidote).

Leaves Usually alternate (rarely distichous; rarely opposite [Cheirodendron] or verticillate), simple, pinnately compound (sometimes two or three times) or palmately compound (leaflets usually articulated), usually pinnately or palmately lobed (sometimes entire), often coriaceous, with ? or supervolute ptyxis. Stipules cauline or petiolar, or one intrapetiolar or absent; lower part of petiole usually sheathing and expanding. Petiole vascular bundles? Venation pinnate or palmate. Stomata anomocytic, paracytic or anisocytic. Cuticular wax crystalloids as irregular platelets and threads. Secretory canals and cavities abundant. Leaf margin and leaflet margins usually serrate (rarely entire); each tooth with broad glandular apex with one main vein and two lateral veins, or with one vein running above tooth.

Inflorescence Terminal or axillary, panicle, thyrsoid etc., or umbellate, spicate or capitate, often compound; partial inflorescences umbels.

Flowers Actinomorphic, small. Usually epigyny (rarely hypogyny or half epigyny). Sepals five, usually minute (sometimes absent), usually with open aestivation, persistent, usually connate. Petals (three to) five to ten (to twelve), with valvate aestivation, caducous, usually free (in, e.g., Osmoxylon connate at base; in, e.g., Boerlagiodendron connate and tubular; in, e.g., ‘Tupidanthus’ [Schefflera] calyptrate); corolla tube initiation early. Nectariferous disc intrastaminal, annular, distally grading into stylopodium.

Androecium Stamens (three to) five to ten (to twelve; rarely twice as many as petals, or numerous; in ‘Tupidanthus’ [Schefflera]calyptratus up to 172 in one whorl; in ‘Plerandra’ [Schefflera] up to c. 500 in four whorls), antesepalous, alternipetalous. Filaments free from each other, from petals and from pistil, inserted outside annular nectariferous disc. Anthers dorsifixed, versatile?, usually tetrasporangiate (rarely octosporangiate), introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–6)-colporate or (2–)3(–6)-colpate, shed as monads, usually tricellular (rarely bicellular) at dispersal. Exine tectate or semitectate, with columellate infratectum, imperforate to finely perforate or reticulate.

Gynoecium Pistil composed of usually two to five (rarely numerous; in ‘Tupidanthus’ [Schefflera] calyptratus up to 132) connate carpels; vascular bundles of ventral carpels consisting of connate bundles of adjacent placentae. Ovary usually inferior (rarely superior or semi-inferior), usually bilocular to quinquelocular (rarely pseudomonomerous or multilocular; in Seemannaralia unilocular). Stylodia two to five, usually free (sometimes connate into single style), inserted on epigynous stylopodium. Stigmas punctate, papillate, usually Wet (sometimes Dry) type. Pistillodium absent.

Ovules Placentation apical. Ovules usually one (rarely two) per carpel, anatropous, pendulous, epitropous, unitegmic, usually crassinucellar (sometimes tenuinucellar). Funicle with short hairs. Integument five to ten cell layers thick. Oburator funicular. Nucellar cap sometimes present. Hypostase sometimes present. Endothelium sometimes present. Megagametophyte usually monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria absent. Embryogenesis onagrad or chenopodiad.

Fruit Usually a drupe with pyrenes of the same number as locules, often dry, with thick-walled lignified mesocarp containing solitary rhomboidal crystals in cell layers immediately outside sclerified bony endocarp (sometimes a berry or syncarp consisting of more or less connate ovaries; rarely [in, e.g., Harmsiopanax with two carpels and carpophore] drupaceous schizocarp with two to five mericarps). “Pseudo-wings”, if present, formed by compression or folding of carpel and composed of both mesocarp and endocarp; vascular bundle present at margin. Vallecular and intrajugal vittae with secretory oil canals and oil cavities usually present.

Seeds Aril absent. Seed coat exotestal. Exotestal cell walls sometimes slightly thickened. Endotesta? Testal cells sometimes with stellate calcium oxalate crystals. Perisperm not developed. Endosperm copious, oily (sometimes ruminate), with petroselinic acid. Embryo small, straight?, well differentiated, without chlorophyll. Cotyledons two. Germination phanerocotylar.

Cytology x = (9) (11) 12 (18)

DNA Mitochondrial gene rpl2 lost. Deletion of 92 bp in plastid gene rpl16. Mitochondrial coxI intron present in Hydrocotyle.

Phytochemistry Flavonols (kaempferol, quercetin), flavonoid sulphates, hydroxycoumarins, oleanolic acid derivatives, dammaranes, sesquiterpene lactones, triterpenoid ethereal oils and resins, caffeic acid, chlorogenic acid, ellagic acid, proanthocyanidins (prodelphinidins), alkaloids, triterpene saponins, acetate derived arthroquinones?, phenantrenes, phenylpropenes (e.g. myristicin), acetophenones, germacrane-like compounds, asarone, syringaresinol, pinoresinol, polyacetylenes (e.g. falcarinone) derived from fatty acids, mostly aliphatic C17-acetylenes, and petroselinic acid (in endosperm) present. Furanocoumarins, tannins and cyanogenic compounds not found. Carbohydrates stored as trisaccharide umbelliferose.

Use Ornamental plants, medicinal plants (Acanthopanax, Aralia, Panax etc.), paper (Tetrapanax papyrifera), timber.

Systematics Araliaceae are sister-group to [Myodocarpaceae+Apiaceae]].

Hydrocotyloideae Link, Handbuch 1: 314. Jan-Aug 1829

3–4/c 260. Trachymene (c 60; Southeast Asia, Malesia, New Guinea, Australia, New Caledonia, Fiji; incl. Uldinia?), Uldinia (1; U. ceratocarpa; southern parts of Australia; in Trachymene?), Hydrocotyle (c 200; cosmopolitan), Neosciadium (1; N. glochidiatum; southwestern Western Australia). – Cosmopolitan, with their highest diversity on the Southern Hemisphere. More or less herbaceous. Nodes 3:3, trilacunar with three leaf traces. Stipules cauline or basal on leaves. Lamina rounded-peltate. Leaf margin crenate. Inflorescence usually terminal (sometimes axillary). Sepals usually absent in Hydrocotyle. Pistil composed of two connate carpels. Integument approx. five cell layers thick (Hydrocotyle). Funicle short. Fruit a schizocarp, laterally compressed. Endocarp sclerified. Carpophore in Trachymene undivided (carpophores not free). n = 6 or higher. Polyploidy frequently occurring in Hydrocotyle. – Hydrocotyloideae may be sister to the remaining Araliaceae, although in some analyses it takes a position as an ingroup in Aralioideae.

Aralioideae Eaton, Bot. Dict., ed. 4: 37. Apr-Mai 1836 [‘Araliaceae’] (under construction)

30–35?/1.830–1.840. Harmsiopanax (3; Malesia), Osmoxylon (c 50; Taiwan, Central and East Malesia, western Pacific islands); Astrotricha (c 20; Australia, with their highest diversity in the southeastern parts); Cheirodendron (6; the Hawaiian Islands, the Marquesas), Raukaua (6; Tasmania, New Zealand, southern Chile, southern Argentina), Motherwellia (1; M. haplosciadea; northeastern Queensland), Cephalaralia (1; C. cephalobotrys; eastern Queensland, eastern New South Wales); Cussonia (c 20; tropical and southern Africa, Madagascar, the Mascarene Islands), Seemannaralia (1; S. gerrardii; South Africa); Aralia (c 70; East and Southeast Asia, Malesia, North America); Polyscias (c 160; tropical regions in the Old World eastwards to New Caledonia, with their largest diversity in Madagascar), Meryta (c 30; eastern Australia, New Caledonia, Vanuatu, New Zealand, Three Kings Islands and Hen and Chickens Islands off New Zealand, Norfolk Island, Pitcairn Island, Society Islands, the Marquesas and other islands in the southwestern Pacific; incl. Pseudopanax?), Pseudopanax (12–20; Tasmania, New Zealand, Chile; in Meryta?), Plerandra (33; New Guinea, the Solomon Islands, Vanuatu, Fiji); Heteropanax (8; southern Asia, China), Fatsia (3; southern Japan, Taiwan), Oplopanax (3; Japan, northwestern North America), Kalopanax (1; K. septemlobus; eastern Siberia, China, the Korean Peninsula, Japan), Metapanax (2; southern China, northern Vietnam), Macropanax (17; Himalayas, China, Southeast Asia, West Malesia), Tetrapanax (1; T. papyrifer; southern China, Taiwan), ’Schefflera’ (c 600?; tropical and subtropical regions on both hemispheres; polyphyletic), Sinopanax (1; S. formosanus; Taiwan; in Schefflera?), Oreopanax (80–150; Mexico, Central America, the West Indies, South America to Brazil and Argentina), Dendropanax (c 90; Southeast Asia, the Malay Peninsula, Central and South America), Merrilliopanax (3; eastern Himalayas, Yunnan), Hedera (12–15; Europe, Macaronesia, the Mediterranean, northwestern Africa, central and southern Asia and eastwards to Japan and Taiwan), Trevesia (7–12; India, Southeast Asia, Malesia; incl. Brassaiopsis?), Brassaiopsis (25–40; Himalayas, China, Thailand, Indochina, Malesia; in Trevesia?), Eleutherococcus (c 40; Himalayas, southeastern Siberia, China, the Korean Peninsula, Japan, Southeast Asia, Malesia to the Philippines). – unplaced Aralioideae Anakasia (1; A. simplicifolia; western New Guinea), Woodburnia (1; W. penduliflora; Burma). – Mainly tropical and subtropical areas, few species in temperate regions. Usually trees, shrubs or lianas (sometimes herbs). Leaves simple, or pinnately or palmately compound. Pistil composed of one to five (to more than 130) connate carpels (when five, then carpels antepetalous; when three, then median carpel abaxial). Stylopodium undivided. Integument approx. ten cell layers thick. Parietal tissue sometimes present. Hypostase sometimes present. Testa multiplictive. Exotestal cells tangentially elongated. – Harmsiopanax, with two carpels and carpophore, may be sister to the remaining Aralioideae, although Plunkett & al. (2004) found it to be nested deeply inside Aralioideae, as sister to Aralia and the Polyscias clade. Osmoxylon may be successive sister to Aralioideae above Harmsiopanax. Astrotricha is possibly also basal, although the support is weak.

Cladogram of Araliaceae based on DNA sequence data (Plunkett & Lowry 2001; Nicolas & Plunkett 2009).

Bayesian inference tree of Araliaceae based on DNA sequence data (Plunkett & al. 2004).

GRISELINIACEAE J. R. Forst. et G. Forst. ex A. Cunn.

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Cunningham in Ann. Nat. Hist. 3: 261. Dec 1839 [’Griselineae’]

Griseliniales (J.R. Forst. et G. Forst. ex Cunn.) Takht. ex Reveal et Doweld in Novon 9: 550. 30 Dec 1999

Genera/species 1/7

Distribution New Zealand, Stewart Island, southeastern Brazil, Paraguay, Chile.

Fossils Unknown.

Habit Dioecious, evergreen trees or shrubs (sometimes lianas or epiphytes).

Vegetative anatomy Phellogen ab initio superficial. Peripheral collenchyma? Vessel elements (solitary) with scalariform perforation plates; lateral pits scalariform or opposite, simple pits. Imperforate tracheary xylem elements ? with bordered pits, septate or non-septate. Wood rays multiseriate, heterocellular. Axial parenchyma usually very rare (apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, often vasicentric) or absent. Sieve tube plastids S type? Nodes 3:3, trilacunar with three leaf-traces, or ≥5:≥5, multilacunar with five or more traces. Prismatic calciumoxalate crystals frequent.

Trichomes Hairs unicellular, with granular, tuberous or verrucate outgrowths.

Leaves Alternate (distichous), simple, entire, coriaceous, often asymmetrical, with conduplicate ptyxis. Stipules absent; leaf base often with petiolar margins and adaxially expanded flange enclosing stem/branch, somewhat sheathing. Petiole vascular bundle transection incurved-arcuate. Venation pinnate? Stomata cyclocytic. Cuticular wax crystalloids? Epidermis with mucilage cells; epidermal cells rhomboidal, small. Mesophyll with sclerenchymatous idioblasts. Leaf margin serrate (often spinose-dentate) or entire.

Inflorescence Terminal or axillary, panicle or raceme. Floral prophylls (bracteoles) absent.

Flowers Actinomorphic, small. Pedicel articulated. Epigyny. Sepals five, with open aestivation, connate, in male flowers minute, in female flowers larger. Petals five, with imbricate aestivation, free (absent in female flowers of Griselinia lucida). Nectary absent. Disc fleshy, quinquangular.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments free from each other and from petals. Anthers dorsifixed, versatile, tetrasporangiate, latrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent?

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, ?-cellular at dispersal. Exine tectate, with ? infratectum, striate.

Gynoecium Pistil composed of usually three (sometimes four) connate carpels (sometimes only two carpels fertile). Ovary inferior, usually unilocular (pseudomonomerous; rarely bilocular); transseptal vascular bundles present. Stylodia usually three (sometimes four), short, free or connate below. Stigmas apical or adaxial, type? Pistillodium absent.

Ovules Placentation apical. Ovule one per carpel, anatropous, pendulous, unitegmic, weakly crassinucellar. Integument ? cell layers thick. Obturator funicular. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit Usually a single-seeded berry.

Seeds Aril absent. Testa multilayered?, with outer two layers (and sometimes especially additional third testal layer) with thickened cell walls. Perisperm not developed. Endosperm copious. Embryo small, straight?, narrowly elongate, chlorophyll? Cotyledons two. Germination?

Cytology n = 18

DNA Mitochondrial gene rpl2 absent?

Phytochemistry Insufficiently known. Route I carbocyclic iridoids (griselinoside), caffeic acid, chlorogenic acid, and petroselinic acid (in endosperm) present. Flavonols? Route II decarboxylated iridoids? Polyacetylenes? Ellagic and gallic acids, tannins, proanthocyanidins, and cyanogenic compounds not found. Umbelliferose?

Use Ornamental plants, timber.

Systematics Griselinia (7; New Zealand, Stewart Island, southeastern Brazil, Paraguay, Chile).

Griselinia is sister to the clade [Pittosporaceae+[Araliaceae+[Myodocarpaceae+Apiaceae]]].

MYODOCARPACEAE Doweld

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Doweld, Tent. Syst. Plant. Vasc.: lii. 23 Dec 2001

Genera/species 2/19

Distribution East Malesia, New Guinea, Melanesia, Queensland, Norfolk Island.

Fossils Unknown.

Habit Usually bisexual, evergreen trees or shrubs.

Vegetative anatomy Phellogen ab initio superficial? Peripheral collenchyma? Vessel elements with simple or scalariform perforation plates; lateral pits alternate, bordered pits? Vestured pits? Imperforate tracheary xylem elements very thick-walled libriform fibres with bordered pits, non-septate (vasicentric tracheids present in Delarbrea). Wood rays multiseriate?, homocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty. Sieve tube plastids S type? Nodes multilacunar? Parenchyma and other tissues usually with schizogenous secretory canals and cavities containing ethereal oils, resins, gum or mucilage? Phelloderm sometimes with sclereids. Crystals?

Trichomes Hairs?

Leaves Alternate (spiral), usually pinnately compound (sometimes simple, entire), with ? ptyxis. Stipules present (Myodocarpus) or absent; petiolar base sheathing. Petiole vascular bundles? Venation usually pinnate, brochidodromous. Stomata? Cuticular wax crystalloids? Leaf margins entire or serrate.

Inflorescence Terminal, panicle (with umbel-like partial inflorescences), or compound umbels.

Flowers Actinomorphic, small. Pedicels articulated. Epigyny. Sepals five, with valvate aestivation, free. Petals five, usually with imbricate (in Delarbrea rarely valvate) aestivation, free, often clawed, apically inflexed. Nectariferous disc intrastaminal, fleshy, present on stylopodium.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments incurved in bud, free from each other and from tepals. Anthers dorsifixed, non-versatile?, tetrasporangiate, introrse?, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains usually tricolporate?, shed as monads, ?-cellular at dispersal. Exine semitectate, with columellate infratectum, reticulate or finely reticulate to scabrate.

Gynoecium Pistil composed of two connate carpels; vascular bundles of ventral carpels adnate to bundles of adjacent placentae. Ovary inferior, bilocular. Stylodia two, free, on a stylopodium. Stigmas non-papillate?, Wet type? Pistillodium absent.

Ovules Placentation axile? Ovule one? per carpel, ?-tropous, unitegmic, tenuinucellar? Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development nuclear? Endosperm haustoria absent? Embryogenesis?

Fruit A drupe (Delarbrea) or schizocarp with free carpophores and two samaroid laterally flattened mericarps (Myodocarpus). Wings (“pseudo-wings”) in Myodocarpus formed by compression or folding of carpel and composed of both mesocarp and endocarp; vascular bundle present at margin. Mesocarp with secretory vesicles. Endocarp reduced, with scattered large circular schizogenous oil ducts (different in morphology from vittae in Apiaceae).

Seeds Seeds in Myodocarpus laterally flattened. Aril absent. Exotestal cells? Endotesta? Perisperm not developed. Endosperm copious, oily. Embryo small, straight?, chlorophyll? Cotyledons two. Germination phanerocotylar?

Cytology n = 12 (Delarbrea)

DNA

Phytochemistry Virtually unknown. Flavonols? Furanocoumarins? Triterpenoid ethereal oils and resins? Sesquiterpene lactones? Polyacetylenes? Petroselinic acid? Umbelliferose?

Use Unknown.

Systematics Myodocarpus (12; New Caledonia), Delarbrea (7; Moluccas, Lesser Sunda Islands, New Guinea, the Solomon Islands, northeastern Queensland, New Caledonia, Vanuatu, Norfolk Island).

Myodocarpaceae are sister to Apiaceae.

PENNANTIACEAE J. Agardh

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Agardh, Theoria Syst. Plant.: 301. Apr-Sep 1858 [‘Pennantieae’]

Pennantiales Doweld, Tent. Syst. Plant. Vasc.: lii. 23 Dec 2001

Genera/species 1/2–4

Distribution Eastern Australia, New Zealand, Norfolk Island.

Fossils Unknown.

Habit Dioecious, usually evergreen trees (Pennantia baylisiana shrub).

Vegetative anatomy Phellogen? Collenchyma poorly developed. Pericyclic envelope present. Vessel elements usually with scalariform (rarely also reticulate) perforation plates; lateral pits opposite and/or scalariform, simple or bordered pits. Imperforate tracheary xylem elements tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, or absent. Tyloses absent. Sieve tube plastids ? type. Nodes 3:3, trilacunar with three leaf traces. Crystals and crystal complexes absent.

Trichomes Hairs unicellular (also uncinate); glandular hairs multicellular with uniseriate stalk.

Leaves Alternate (spiral), simple, entire, often coriaceous, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection annular; petiole often with wing bundles and sometimes with medullary bundle. Venation pinnate. Stomata paracytic. Cuticular wax crystalloids? Domatia as pockets or hair tufts, or absent. Foliar sclereids absent. Leaf margin serrate, crenate or entire.

Inflorescence Usually terminal (sometimes axillary), panicle.

Flowers Actinomorphic, small. Pedicel articulated. Hypogyny. Sepals four or five, with open? aestivation, very small, free. Petals five, with valvate aestivation, connate at base, inflexed at apex. Nectary absent. Disc absent.

Androecium Stamens four or five?, haplostemonous, antesepalous, alternipetalous. Filaments free from each other, folded in bud, sometimes adnate to corolla (epipetalous). Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Female flowers with or without staminodia.

Pollen grains Microsporogenesis simultaneous? Pollen grains?, shed as monads, ?-cellular at dispersal. Exine?, with ? infratectum, sculpturing?

Gynoecium Pistil composed of two or three connate carpels. Ovary superior, unilocular?; transseptal vascular bundles present? Stylodia three, very short (when stigmas punctate), or styles absent. Stigmas three, either punctate or hippocrepomorphic forming sinuate ring, papillate, type? Male flowers with pistillodium.

Ovules Placentation apical. Ovules one per carpel, anatropous, unitegmic, tenuinucellar or thinly crassinucellar. Integument ? cell layers thick, vascularized. Megagametophyte monosporous, Polygonum type. Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A drupe with inner cells transversely elongate.

Seeds Aril absent. Testa thin. Exotestal cells? Endotesta? Perisperm? Endosperm copious, oily?, chlorophyll? Embryo minute, straight? Cotyledons two. Germination?

Cytology n = 25

DNA Mitochondrial gene rpl2 absent?

Phytochemistry Unknown. Iridoids? Umbelliferose?

Use Timber.

Systematics Pennantia (2–4; eastern Australia, New Zealand, including the Three Kings Islands, Norfolk Island).

Pennantia is sister to the remaining Araliales.

PITTOSPORACEAE R. Br.

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Brown in M. Flinders, Voy. Terra Austral. 2: 542. 19 Jul 1814 [’Pittosporeae’], nom. cons.

Pittosporales Link, Handbuch 2: 222. 4-11 Jul 1829 [’Pittosporeae’]

Genera/species 8/c 200

Distribution Madeira, Africa south of Sahara, Madagascar, Yemen, India, Sri Lanka, eastern Himalaya, East Asia to the Korean peninsula and southern Japan, Southeast Asia, Malesia, New Guinea, Melanesia, Australia, Tasmania, New Zealand, islands in the Pacific and the Indian Ocean, with their largest diversity in Australia.

Fossils Unknown.

Habit Usually bisexual (sometimes polygamomonoecious, dioecious, or gynodioecious), evergreen trees, shrubs or lianas (sometimes spiny). Some species are xerophytic. Many species are aromatic.

Vegetative anatomy Phellogen ab initio superficial. Peripheral collenchyma? Young stem with vascular cylinder (in Pittosporum septated). Vessel elements usually with simple (in one species of Pittosporum scalariform) perforation plates (in primary xylem with scalariform perforation plates only); lateral pits alternate, bordered pits. Vestured pits? Imperforate tracheary xylem elements fibre tracheids and/or libriform fibres with simple or bordered pits, septate to non-septate (also vasicentric tracheids). Wood rays multiseriate, heterocellular (almost homocellular). Axial parenchyma paratracheal scanty, often vasicentric, or absent. Sieve tube plastids Ss type. Nodes usually 3:3, trilacunar with three leaf traces (sometimes 1:3, unilacunar with three traces). Parenchyma and other tissues (including pericycles of stem, secondary cortex and secondary phloem of older stems, roots and leaves) with schizogenous secretory canals and cavities with ethereal oils? and/or resins and/or gum-like substances? Prismatic calciumoxalate crystals often frequent.

Trichomes Hairs unicellular or multicellular, uniseriate or branched, short-stalked T-shaped with two or three basal cells and long terminal cell (terminal cell vertically or transversely elongate); glandular hairs clavate or absent.

Leaves Alternate (spiral), simple (leaves of seedlings in Pittosporum sometimes pinnately compound), usually entire (rarely lobate), often coriaceous, with supervolute to curved ptyxis. Stipules absent; leaf base narrow to sheathing and decurrent. Petiole vascular bundle transection arcuate. Venation pinnate. Stomata usually paracytic (rarely cyclocytic). Cuticular wax crystalloids as rodlets. Schizogenous secretory canals and cavities (around phloem) with resin. Mesophyll cells with calciumoxalate usually as druses. Leaf margin usually entire (rarely serrate or sinuate).

Inflorescence Terminal or axillary, corymb, panicle, thyrse etc., or flowers solitary. Partial inflorescences not umbels.

Flowers Usually actinomorphic (sometimes slightly obliquely zygomorphic). Hypogyny (probably secondary). Sepals five, with imbricate aestivation, caducous, usually free (sometimes connate at base); median sepal adaxial. Petals five, with imbricate aestivation, usually connate at base. Nectariferous disc intrastaminal.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments free from each other or somewhat connate at base, free from petals. Anthers usually dorsifixed (sometimes basifixed), non-versatile, tetrasporangiate, introrse, usually longicidal (dehiscing by longitudinal or sometimes short slits; in Cheiranthera poricidal, with apical pores). Placentoid present. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenes simultaneous. Pollen grains 3(–4)-colporate, shed as monads, usually tricellular (sometimes bicellular) at dispersal. Exine tectate, with columellate? infratectum, imperforate to finely perforate.

Gynoecium Pistil composed of two (to five) connate carpels. Ovary superior (probably secondarily), usually unilocular (rarely completely or incompletely bilocular to quinquelocular), with lateral nectaries. Style one, entire. Stigma capitate or somewhat lobate, non-papillate, Wet type. Pistillodium absent.

Ovules Placentation parietal (when ovary unilocular) or axile (when ovary multilocular); ovules inserted above basal zone of separate loculi (comparable to Araliaceae and Apiaceae). Ovules several to numerous per carpel, anatropous or somewhat campylotropous, horizontal or ascending, usually epitropous (sometimes apotropous), unitegmic, tenuinucellar. Integument ? cell layers thick. Funicular obturator probably absent. Endothelium absent. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis asterad.

Fruit A loculicidal (sometimes also septicidal) capsule or a berry. Seeds surrounded by resinous, juicy and viscid tissue.

Seeds Aril present or absent. Seed coat exotestal, fleshy. Testa multilayered. Exotestal cell walls thickened, non-lignified. Endotesta? Perisperm not developed. Endosperm copious, oily. Embryo small, straight, well differentiated, without chlorophyll. Cotyledons usually two (in Pittosporum sometimes up to five). Germination phanerocotylar.

Cytology x = 12

DNA Plastid gene infA lost/defunct (Pittosporum). Mitochondrial gene rpl2 lost?

Phytochemistry Furanocoumarins, hydroxycoumarins, oleanolic acid derivatives, non-hydrolyzable tannins, triterpenoid ethereal oils and resins, caffeic acid, alkaloids, triterpene saponins, polyacetate derived arthroquinones, C20- and C22-fatty acids, polyacetylenes (e.g. falcarinone) derived from fatty acids, mostly aliphatic C17-acetylenes, and chinic acid present. Ellagic acid, proanthocyanidins, cyanogenic compounds and petroselinic acid not found. Flavonols (kaempferol, quercetin)? Carbohydrates stored as trisaccharide umbelliferose. Aluminium accumulated.

Use Ornamental plants, carpentries.

Systematics Bursaria (7; eastern, southeastern and southwestern Australia, Tasmania), Auranticarpa (6; northern Australia to eastern New South Wales), Pittosporum (c 200; Madeira [Pittosporum coriaceum], tropical and southern Africa, Madagascar, the Arabian Peninsula, southern and eastern Asia, Australia, Tasmania, New Zealand, Pacific islands incl. the Hawaiian Islands), Hymenosporum (1; H. flavum; New Guinea, eastern Queensland, northeastern New South Wales), Bentleya (2; southwestern Western Australia), Billardiera (c 25; Australia, Tasmania), Marianthus (c 15; southwestern and southeastern Australia, Tasmania), Cheiranthera (5; southwestern and southeastern Australia).

Pittosporaceae are sister-group to [Myodocarpaceae+Apiaceae].

Bursaria may be sister to the remaining Pittosporaceae, although the support is weak and the basal clade usually a trichotomy.

Cladogram of Pittosporaceae based on morphological data (Cayzer & al. 2004).

TORRICELLIACEAE (Wang) H. H. Hu

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Hu in Bull. Fan Mem. Inst. Biol., Bot., ser. 5, 5: 311. 1934

Melanophyllaceae Takht. ex Airy Shaw in Kew Bull. 26: 492. 1972; Aralidiaceae Philipson et B. C. Stone in Taxon 29: 402. 25 Aug 1980; Aralidiales Takht. ex Reveal in Novon 2: 238. 13 Oct 1992; Torricelliales Takht. ex Reveal et Doweld in Novon 9: 550. 30 Dec 1999

Genera/species 3/12

Distribution Madagascar, eastern Himalaya to northern Burma and China, southern Thailand, West Malesia.

Fossils A fossilized fruit attributed to Torricellia has been described from the Eocene of North America. Numerous additional macrofossils have been assigned to Torricelliaceae.

Habit Bisexual (Melanophylla), monoecious (Torricellia) or dioecious (Aralidium), evergreen or deciduous trees or shrubs.

Vegetative anatomy Phellogen ab initio superficial (Aralidium). Primary cork tissue in Torricellia with collenchyma. Cortical vascular bundles present in Aralidium. Endodermis absent (Melanophylla). Vessel elements with simple (Melanophylla, Torricellia) or scalariform (Aralidium, Melanophylla) perforation plates; lateral pits in Aralidium opposite or scalariform, in Melanophylla alternate or scalariform, in Torricellia alternate or opposite, simple pits. Imperforate tracheary xylem elements libriform fibres with simple (Aralidium) or bordered (Melanophylla, Torricellia) pits, septate or non-septate. Wood rays multiseriate, heterocellular. Axial parenchyma apotracheal (Melanophylla, Torricellia) or paratracheal scanty, vasicentric (Aralidium, Melanophylla). Tyloses frequent. Sieve tube plastids S type? Nodes ≥5:≥5, multilacunar with five or more leaf traces (Aralidium, Torricellia). Secretory canals absent. Medulla in Aralidium heterogeneous, with groups of sclereids. Calciumoxalate as crystal sand present in Aralidium.

Trichomes Hairs unicellular or multicellular, uniseriate (absent in Melanophylla); glandular hairs in Aralidium multicellular asymmetrical, in Torricellia tricellular or quadricellular.

Leaves Alternate (spiral), simple, entire (Melanophylla), pinnately (Aralidium) or palmately (Torricellia) lobed, with ? ptyxis. Stipules absent (Aralidium, Torricellia) or present as glandular hairs (Melanophylla); petiole sheathing at base. Ligule sometimes present. Petiole vascular bundles scattered (Aralidium). Leaf base with petiolar flange (and sometimes small adaxial flange), often enclosing branch. Venation pinnate (Aralidium, Melanophylla) or palmate (Torricellia). Stomata anomocytic (Melanophylla, Torricellia) or anisocytic (Aralidium). Cuticular wax crystalloids? Mucilage cells present. Leaf margin usually serrate and/or sinuate (sometimes entire).

Inflorescence Terminal or axillary, usually panicle (sometimes thyrsoid; Aralidium, Torricellia, Melanophylla) or raceme (Melanophylla).

Flowers Actinomorphic, small. Pedicel usually articulated (in female flowers of Torricellia inarticulated). Epigyny. Sepals three to five, with imbricate or open aestivation (Torricellia), free (Aralidium) or connate (Torricellia, Melanophylla). Petals five, with imbricate (Aralidium, Melanophylla) or induplicate-valvate (male flowers in Torricellia) aestivation, free (absent in female flowers of Torricellia). Nectaries in Melanophylla present on stout stylar discs. Disc absent in Torricellia; vascularized intrastaminal nectariferous disc present in female flowers of Aralidium as usually three (rarely four) convex stylopodia.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments short, thin, free from each other and from tepals. Anthers basifixed (Melanophylla, Torricellia) or dorsifixed (Aralidium, Melanophylla), versatile?, tetrasporangiate, latrorse to introrse (in Torricellia latero-introrse), longicidal (dehiscing by longitudinal slits). Tapetum secretory? Female flowers in Aralidium with staminodia.

Pollen grains Microsporogenesis simultaneous? Pollen grains 3(–4)-colporate, usually shed as monads (in Melanophylla usually as tetrads), ?-cellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate, reticulate, rugulate or smooth.

Gynoecium Pistil composed of usually three (sometimes two or four) connate carpels; transseptal vascular bundles present. Ovary inferior, trilocular (sometimes bilocular or quadrilocular; Torricellia, Melanophylla) or secondarily unilocular (pseudomonomerous, only one locule fertile in Melanophylla and Aralidium). Stylodia usually three (sometimes two or four), subulate, erect or reflexed, free (in Aralidium on stylopodia), or absent. Stigmas linear or punctate (in Torricellia bilobate, persistent), type? Male flowers in Torricellia with pistillodium.

Ovules Placentation apical (in Melanophylla axile to apical). Ovules one per carpel, anatropous, pendulous, apotropous, unitegmic, tenuinucellar. Integument ? cell layers thick (massive in Aralidium). Funicular obturator present in Torricellia. Megagametophyte monosporous, Polygonum type. Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Fruit A single-seeded drupe with one fertile locule and two to four degenerated sterile locules (often with persistent styles and stigmas). Endocarp with sclereids.

Seeds Aril present (in Torricellia formed by funicular obturator). Testa very thin (in Aralidium vascularized; in Torricellia somewhat sclerified). Exotestal cells with scalariform thickened walls, non-lignified (Melanophylla). Endotesta? Perisperm absent (Aralidium). Endosperm copious, fleshy, usually ruminate. Embryo small, well differentiated, straight (Aralidium, Melanophylla) or curved and thin (Torricellia), chlorophyll? Cotyledons two. Germination?

Cytology n = 12 (Torricellia), 20±2

DNA Mitochondrial gene rpl2 absent?

Phytochemistry Insufficiently known. Flavonols (kaempferol, rare), Route I iridoid glucosides (griselinoside in Torricellia, aralioside and verbascosides in Aralidium), ellagic acid, tannins (Torricellia), and C11 polyacetylenes with chiral centre (Torricellia) present. Route II iridoids? Proanthocyanidins not found.

Use Unknown.

Systematics Torricellia (3; western Nepal to Bhutan, northern Burma, continental China), Aralidium (1; A. pinnatifidum; southern Thailand, the Malay Peninsula, Sumatra, Borneo), Melanophylla (8; Madagascar).

Torricelliaceae are sister to the clade [Griseliniaceae+[Pittosporaceae+[Araliaceae+[Myodocarpaceae+Apiaceae]]]].

Aralidium is sister to [Melanophylla+Torricellia] (Soltis & al. 2011).


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