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Araliaceae
From Wikipedia, the free encyclopedia Araliaceae
Temporal range: Eocene–0PreЄЄOSDCPTJKPgN | | Aralia elata | Scientific classification | Kingdom: | Plantae | (unranked): | Angiosperms | (unranked): | Eudicots | (unranked): | Asterids | Order: | Apiales | Family: | Araliaceae
Juss.[1] | Subfamilies and genera | * See text | Synonyms | * Botryodendraceae J.Agardh * Hydrocotylaceae (Drude) Hyl., nom. cons. |
Araliaceae is a family of flowering plants, also known as the Aralia family (after its type genus Aralia) or ivy family. The family includes 254 species of trees, shrubs, lianas and perennial herbaceous plants in two subfamilies. Species usually bear pinnately or palmatelycompound leaves, and usually have small flowers produced in large panicles.
Contents
[hide] * 1 Overview * 2 Subfamilies and genera * 3 See also * 4 References and external links
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Overview[edit]
The family from tropical area origin is present in cooler climates, too. They are found in the Americas, Eurasia, Africa, Australia, New Zealand, New Caledonia and Pacific islands. Araliaceae bear essential oils, or without essential oils can be resinous and heterophyllous. It presents many shapes, includes some trees and ivies as the angelica tree (devil's walking-stick, Aralia spinosa), the devil's club (Oplopanax horridus), Hedera spp., including Hedera helix and herbs as ginseng Panax spp., a native of Korea and used as medical herb. Leaves are simple, or compound, sometimes lauroid (resembling Laurus) or peltate, or not peltate; when compound, ternate, or pinnate, or palmate.
The systematics of Araliaceae are currently under study, and taxonomic changes and novelties are to be expected. Endemic Araliaceae are found in the pluvial montane forest, very humid montane, and humid lowland river forest regions. They are present, too, in laurel forest, cloud forest and warm, humid habitats. The family is closely related to Apiaceae andPittosporaceae, and the boundaries between these families and other members of Apialesare still uncertain. Some recent systems included Araliaceae in an expanded Apiaceae, but this has not been widely followed. Molecular phylogenies[2] suggest at least some of the genera traditionally included in Apiaceae as subfamily Hydrocotyloideae appear to be more closely related to Araliaceae, and the inclusion of Hydrocotyle and Trachymene in Araliaceae has been recommended.[3]
The generic level classification of Araliaceae has been unstable; in particular, numerous genera have been synonymized underSchefflera. Recent molecular phylogenies[4] have shown this large pantropical genus is polyphyletic and some believe it should be divided again into several genera, though these would probably not correspond with the previously recognized genera.
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Subfamilies and genera[edit] Subfamily Aralioideae * Anakasia * Apiopetalum * Aralia * Arthrophyllum * Astrotricha * Boninofatsia * Brassaiopsis * Cephalaralia * Cheirodendron * Cromapanax * Cuphocarpus * Cussonia * Dendropanax * Eleutherococcus * Fatshedera * Fatsia * Gamblea * Gastonia * Harmsiopanax * Hedera * Heteropanax * Hunaniopanax * Kalopanax * Mackinlaya * Macropanax * Megalopanax * Merrilliopanax * Meryta | * Metapanax * Motherwellia * Munroidendron * Oplopanax * Oreopanax * Osmoxylon * Panax * †Paleopanax[5] * Polyscias * Pseudopanax * Pseudosciadium * Raukaua * Reynoldsia * Schefflera * Sciadodendron * Seemannaralia * Sinopanax * Stilbocarpa * Tetrapanax * Tetraplasandra * Trevesia * WoodburniaSubfamily Hydrocotyloideae * Azorella * Centella * Hydrocotyle * Xanthosia | *
Schefflera arboricola *
(Oplopanax horridus) *
Eleutherococcus sieboldianus *
Hedera helix *
Aralia spinosa *
Osmoxylon lineare

Araliaceae
The Araliaceae are mostly tropical shrubs and trees comprising about 70 genera and 700 species. The leaves are alternate or rarely opposite, palmately or pinnately compound or more than once compound or rarely simple; stipules are usually present and liguliform or adnate to the petiole and sheathing. The flowers are actinomorphic and most frequently unisexual, often in heads or umbels. The perianth is biseriate but the calyx is reduced to usually 5 minute teeth or a seamlike rim adnate to the ovary. The corolla consists mostly of 5-10 usually more or less distinct, usually valvate petals arising from a nectary disk on the summit of the ovary. The stamens are distinct, usually as many as and alternating with the petals. The gynoecium consists of a single compound pistil of 2-15 carpels, an equal number of styles or these connate into one style, and an inferior ovary with 2-15 locules, each bearing a single pendulous, axile ovule. An epigynous nectary disk is generally confluent with the enlarged stylar base or stylopodium. The fruit is a berry or drupe that sometimes splits into one-seeded segments.
Each "thumbnail" image below is linked to a larger photograph.

| | | Arthrophyllum sp. Note the palmately compound leaf, 5-merous flowers, umbellate inflorescence and the calyx that forms a rim with minute teeth around the summit of the inferior ovary. | | Cheirodendron platyphyllum, 'olapa. This Hawaiian endemic species has leaves (and leaflets) that quake like aspen (for the same reason, i.e. flattened stalks). | | | | Munroidendron racemosum. This is a rare Hawaiian endemic genus that has the flowers in racemes instead of the usual umbels found in the family. It also has numerous stamens and irregularly connate petals. | | Oplopanax horridum, devil's walking cane, devil's club, Hackleman Old Growth Trail, Cascades, OR, July 2003. | | | | Reynoldsia sandwicensis, 'ohe. In this endemic Hawaiian species the petals are often connate in pairs or threes, giving the appearance of commonly 4 or 5 petals. There are usually about 10 stamens. | | | | Schefflera actinophylla, octopus tree. This ornamental species from Australia has flowers with about a dozen perianth segments and stamens. The petals fall away that is, are caducous as the stamens expand. A conspicuous disk is present and a style is lacking. | | | | | Tetraplasandra oahuensis, 'ohe mauka. Note the pinnate leaves, umbellate inflorescence, and flowers with five and six petals and alternating stamens; the inferior ovary and nectary disk are also evident in this this endemic Hawaiian species. | | Tetraplasandra waimeae, 'ohe kiko'ola, endemic genus. |

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Classification for Kingdom Plantae Down to Family AraliaceaeClick on names to expand them, and on P for PLANTS profiles. | | Up to the Kingdom | Kingdom Plantae – Plants | Subkingdom Tracheobionta – Vascular plants | Superdivision Spermatophyta – Seed plants | Division Magnoliophyta – Flowering plants | Class Magnoliopsida – Dicotyledons | Subclass Rosidae | Order Apiales | Family Araliaceae – Ginseng family | | Contains 18 Genera and 47 accepted taxa overall | | | | | | | | | | | | | | | | | | | | | | | | | | Down one level | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | Genus Aralia L. – spikenard P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Cheirodendron Nutt. ex Seem. – cheirodendron P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Dendropanax Decne. & Planch. – dendropanax P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Eleutherococcus Maxim. – ginseng P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Fatsia Decne. & Planch. – fatsia P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Hedera L. – ivy P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Kalopanax Miq. – castor aralia P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Meryta J.R. Forst. & G. Forst. P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Munroidendron Sherff – munroidendron P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Oplopanax (Torr. & A. Gray) Miq. – oplopanax P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Osmoxylon Miq. P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Panax L. – ginseng P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Polyscias J.R. Forst. & G. Forst. – aralia P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Pseudopanax K. Koch – pseudopanax P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Reynoldsia A. Gray – reynoldsia P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Schefflera J.R. Forst. & G. Forst. – schefflera P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Tetrapanax (K. Koch) K. Koch – tetrapanax P | | | | | | | | | | | | | | | | | | | | | | | | | Genus Tetraplasandra A. Gray – tetraplasandra P | | | | | | | | | | | | | | | | | | | | | | | | | | |

| The families of flowering plants | |
L. Watson and M. J. Dallwitz
Araliaceae Juss.
Including Botryodendraceae J.G. Agardh, Myodocarpaceae Doweld; excluding currently Umbelliferae-Hydrocotyloideae.
Habit and leaf form. Trees (mostly, usually of moderate size but sometimes very large, with Peekeliopanax reaching 40 m high), or ‘arborescent’, or shrubs (including some woody epiphytes), or lianas, or herbs (Panax, Stilbocarpa, some species of Aralia, etc.); non-laticiferous and without coloured juice; bearing essential oils, or without essential oils; resinous. ‘Normal’ plants (nearly always), or switch-plants (e.g., with the linear leaves of Lilaeopsis and Oxypolis interpretable as highly modified pinnately compound leaves, with nodal appendages representing pinnae transformed into hydathodes). Plants autotrophic. Perennial; without conspicuous aggregations of leaves. Self supporting, or epiphytic, or climbing; when climbing stem twiners, or root climbers. Pachycaul (nearly always, mostly large-leaved and thick-stemmed), or leptocaul (e.g., in Pseudopanax, in which long- and short-shoots are distinguishable). Heterophyllous (sometimes, e.g. Hedera helix, where progression from lobed to entire leaves reflects irreversible shoot maturation), or not heterophyllous. Leaves medium-sized to very large (to over 3 m inAralia), or small (rarely, but only 1–2 cm in Pseudopanax anomalum); alternate (nearly always), or opposite (Cheirodendron, Eremopanax), or whorled (Panax); spiral (mostly), or distichous (rarely), or four-ranked (rarely); commonly leathery; petiolate (usually), or subsessile; more or less sheathing (usually), or non-sheathing. Leaf sheaths with free margins. Leaves gland-dotted, or not gland-dotted; aromatic, or without marked odour; simple (mostly), or compound; not peltate (usually), or peltate (some Harmsiopanax spp.); when compound, ternate, or pinnate, or palmate, or multiply compound. Lamina when simple, dissected (usually), or entire; pinnatifid, or palmatifid; pinnately veined, or palmately veined. Leaves stipulate, or exstipulate. Stipules when present, intrapetiolar (often adnate to and scarcely distinguishable from the base of the petiole). Vegetative buds scaly. Leaf development not ‘graminaceous’; leaves becoming compound from primordial lobes.
Leaf anatomy. The leaf lamina dorsiventral (usually), or bifacial to centric. Stomata present; usually mainly confined to one surface (mostly abaxial); paracytic (usually), or anomocytic. Hairs present (not always numerous, but of diverse types, with shaggy, 2-armed, tufted, stellate, and peltate forms recorded). Complex hairs present (usually), or absent; commonly stellate. Adaxial hypodermis present (very commonly), or absent. Lamina or more often canals with secretory cavities. Secretory cavities presumably containing resin, or containing oil, containing mucilage, containing resin, and containing latex (? - cf. those of the stems); schizogenous. The mesophyll not containing mucilage cells (seemingly lacking secretory cells of any sort). Minor leaf veins without phloem transfer cells (Aralia, Hedera).
Axial (stem, wood) anatomy. Secretory cavities or more often canals universally present; with resin (presumably), or with oil, with resin, with mucilage, and with latex (i.e., with imprecise references to oily, resinous, gummy and occasionally milky contents). Cork cambium present; initially superficial. Nodes penta-lacunar to multilacunar (mostly), or tri-lacunar (rarely). Primary vascular tissue generally comprising a ring of bundles (at first, separated by wide rays), or in a cylinder, without separate bundles (subsequently), or comprising two or more rings of bundles, or consisting of scattered bundles (the conventional ring often accompanied by additional cortical and/or medullary circles of bundles, and occasionally by scattered medullary bundles); collateral. Internal phloem absent. Cortical bundles present (commonly), or absent. Medullary bundles present (often), or absent. Secondary thickening usually developing from a conventional cambial ring (? - no reference to anomalous secondary thickening having been found).
The wood ring porous to diffuse porous. The vessels small to medium; variously radially paired, in radial multiples, clustered, and in tangential arcs. The vessel end-walls simple (usually), or scalariform (often with few bars), or scalariform and simple. The vessels without vestured pits; with spiral thickening, or without spiral thickening. The axial xylem without tracheids; without fibre tracheids; with libriform fibres; usually including septate fibres (especially around the vessels). The fibres without spiral thickening. The parenchyma usually paratracheal (only, often very sparse). ‘Included’ phloem absent. The wood not storied. Tyloses present, or absent.
Reproductive type, pollination. Plants hermaphrodite, or monoecious, or andromonoecious, or gynomonoecious, or dioecious, or polygamomonoecious.
Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in spikes, in heads, and in umbels (with umbels less prevalent than in the Umbelliferae). The ultimate inflorescence units cymose (mostly), or racemose (represented in probably no more than 10 genera). Inflorescencesterminal, or axillary, or leaf-opposed (though appearing lateral, rarely - e.g. in Mackinlaya, and sometimes genuinely lateral in Aralia, Schefflera and Stilbocarpa), or epiphyllous (rarely); a few genera exhibiting racemes, most with umbels or heads, often massed into compound inflorescences. Flowers calyptrate (rarely), or not calyptrate; usually more or less 5 merous; cyclic.
Perianth with distinct calyx and corolla, or petaline; (6–)10(–24); 1 whorled (only in Meryta, where the calyx is entirely lacking), or 2 whorled; when two-whorled, isomerous, or anisomerous. Calyx when present, 3–5(–12); 1 whorled; when present, polysepalous, or gamosepalous; entire, or lobulate, or blunt-lobed, or toothed (sometimes reduced to small teeth or a mere rim); often open in bud. Corolla (3–)5(–13) (the interpretation sometimes complicated by partially divided or lobed segments); 1 whorled; commonly alternating with the calyx (most genera having five calyx components alternating with five petals, but numerous exceptions occur); polypetalous, or partially gamopetalous, or gamopetalous (and sometimes connate at the base); calyptrate (rarely), or not calyptrate; imbricate (in the Aralieae), or valvate; regular; typically rather fleshy, or not fleshy. Petals clawed (Mackinleyeae), or sessile (usually with broad bases inserted around the whole circumference of the upper part of the ovary).
Androecium (3–)5(–12), or 10–100 (the same number as the corolla members, twice the number, or ‘many’). Androecial members free of the perianth; all equal; free of one another. Androecium exclusively of fertile stamens. Stamens variously (3–)5(–12), or 10–100; isomerous with the perianth (usually), or diplostemonous, or polystemonous; alternating with the corolla members (usually, when equalling them in number), or opposite the corolla members;inflexed in bud; filantherous (the filaments usually fleshy and short). Anthers dorsifixed; not becoming inverted during development; dehiscing via longitudinal slits; introrse; tetrasporangiate (nearly always, but occasionally appearing bisporangiate by early fusion dring development), or multisporangiate (8 inOctotheca and Dizygotheca). Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral. Anther wall initially with one middle layer, or initially with more than one middle layer. Tapetum glandular. Pollen grains aperturate; 3 aperturate (mostly), or 2 aperturate, or 4 aperturate, or 6 aperturate; colporate (mostly), or colpate, or rugate; 3-celled.
Gynoecium (1–)2–5(–100) carpelled (about 20 in species of Gastonia, Plerandra and Reynoldsia, up to 100 in Tupidanthus). Carpels reduced in number relative to the perianth to increased in number relative to the perianth. The pistil (1–)2–5(–100) celled. Gynoecium syncarpous (nearly always), or monomerous (rarely, at least ostensibly so); of one carpel (in Diplopanax, and ostensibly so in Arthrophyllum and Eremopanax), or synovarious to eu-syncarpous (nearly always, the styles often constituting a solid or sometimes hollow stylopodium); partly inferior to inferior, or superior (recorded only in two species of Tetraplasandra). Carpel when monomeric, i.e. rarely, ostensibly 1 ovuled (but with a second abortive one). Ovary 1–100 locular. Locules without ‘false septa’. Epigynous disk present (with a nectariferous disk between the stylopodium and the stamens). Gynoecium more or less stylate. Styles variously 1–100; when two or more, free, or partially joined; apical. Stigmas usually present, as a frequently double stigmatic crest capping the stylopodium; wet type, or dry type; papillate; Group II type and Group III type. Placentation when unilocular, i.e. rarely, parietal to apical; when bi- or plurilocular, i.e. nearly always, axile to apical. Ovules in the single cavity 1–2 (with one abortive); 1 per locule, or 1–2 per locule (usually with a second abortive one); pendulous; epitropous; with ventral raphe; anatropous;unitegmic; crassinucellate (mostly), or tenuinucellate. Endothelium differentiated. Polar nuclei fusing prior to fertilization. Antipodal cells formed, or not formed (then the three nuclei degenerating early); when formed, 3; not proliferating. Synergids pear-shaped. Hypostase present, or absent. Endosperm formation nuclear.
Fruit fleshy, or non-fleshy; indehiscent, or a schizocarp (then cf. Umbelliferae). Mericarps when schizocarpic, 2–5(–100) (?). Fruit when non-schizocarpic, a berry, or a drupe. The drupes with separable pyrenes, or with one stone (as many pyrenes as the locules). Gynoecia of adjoining flowers combining to form a multiple fruit, or not forming a multiple fruit. Seeds endospermic. Endosperm ruminate (e.g. Hedera), or not ruminate; oily. Cotyledons 2. Embryo achlorophyllous (2/3).
Seedling. Germination phanerocotylar.
Physiology, phytochemistry. C3. C3 physiology recorded directly in Hedera. Sugars transported as sucrose (Brassaia, Silibertia). Inulin not found (umbelliferose recorded). Not cyanogenic. Polyacetylenes recorded (falcarinone). Alkaloids present, or absent. Arbutin absent. Iridoids not detected (in any of the members screened, discounting a spurious record for Hedera and with Aralidium and Diplopanax excluded). Saponins/sapogenins present. Proanthocyanidins absent. Flavonols present, or absent; quercetin, or kaempferol and quercetin. Ellagic acid absent (5 species, 5 genera). Aluminium accumulation not found. Sieve-tube plastids S-type.
Geography, cytology. Temperate (a few), or sub-tropical to tropical (mainly). Widespread, but especially Indomalaya and tropical America. X = 11, 12(+).
Taxonomy. Subclass Dicotyledonae; Tenuinucelli (or borderline, but uncertainties may reflect a few misplaced genera). Dahlgren’s Superorder Araliiflorae; Araliales. Cronquist’s Subclass Rosidae; Apiales. APG 3 core angiosperms; core eudicot; Superorder Asteranae; campanulid; Order Apiales.
Species 700. Genera 49; Anakasia, Apiopetalum, Aralia, Arthrophyllum, Astrotricha, Boninofatsia, Brassaiopsis, Cephalaralia, Cheirodendron,Cromapanax, Cuphocarpus, Cussonia, Delarbrea, Dendropanax, Eleutherococcus, Fatsia, Gamblea, Gastonia, Harmsiopanax, Hedera, Heteropanax,Hunaniopanax, Kalopanax, Mackinlaya, Macropanax, Megalopanax, Merrilliopanax, Meryta, Motherwellia, Munroiodendron, Myodocarpus, Oplopanax,Oreopanax, Osmoxylon, Panax, Pentapanax, Polyscias, Pseudopanax, Pseudosciadium, Reynoldsia, Schefflera, Sciadodendrom, Seemannaralia,Sinopanax, Stilbocarpa, Tetrapanax, Tetraplasandra, Trevesia, Woodburnia. Diplopanax has been transferred to the cornaceous alliance.
General remarks. This family exemplifies the well known difficulties in distributing certain Dicot families not only between Dahlgren’s Araliiflorae and Corniflorae, but also between the higher level groupings Crassinucelli and Tenuinucelli; this despite the fact that the latter evidently represent a major divergence in the Dicot line of descent (cf.Young and Watson 1970, Chase et al. 1993). In terms of the data compiled for this package, other than two ‘esoteric characters’ reflecting limited sampling (wet/dry stigmas, occurrence of septate wood fibres), there are no absolute differences between Araliaceae and Umbelliferae: they are distinguishable only in having the states of overlapping characters expressed in different proportions. This does not mean of course that the families should be merged, because the names permit useful predictive generalization. On the other hand, it does support S.M. Walters’s (1960) entertaining contention regarding the historical origins of useful family circumscriptions; viz., that that if formal taxonomy had originated in the southern hemisphere, the Umbelliferae would now be a subfamily of the Araliaceae.
Economic uses, etc. Some cultivated ornamentals, including notable houseplants, e.g. Hedera, Aralia, Polyscias, Schefflera, Fatsia. Ginseng roots from Panax quinquefolius, Chinese rice paper from the pith of Tetrapanax papyriferus.
Illustrations. • Technical details, from Thonner (Cussonia). • Technical details (Hedera, Reynoldsia). • Technical details (Hedera helix: Lindley). • Technical details (Aralia edulis, Gastonia). • Hedera helix (B. Ent.). • Trevesia palmata: Bot. Reg. 894, 1825. • Leaf hairs (Solereder, 1908).
Quotations
Sleep thou, and I will wind thee in my arms,
. . . . the female ivy so
Enrings the barky fingers of the elm
(‘Midsummer Night’s Dream’, iv., 1)
Oh, falsely they accuse me,
Who say I seek to check
The growing sapling’s flourishing; —
I better love to deck
The dead or dying branches
With all my living leaves.
’Tis for the old and wither’d tree,
The Ivy garlands weaves.
Calder Campbell, quoted by Ann Pratt 1857)