Araliaceae

English ivy Hedera helix, copyright Bengt-re.

Belongs within: Apiales.
Contains: Pseudopanax, Polyscias, Osmoxylon, Hydrocotyle, Trachymene, Panax, Schefflera, Aralia.

The Araliaceae are a family of plants found worldwide, with diversity particularly high in the tropics. The majority of Araliaceae are shrubs and trees, though the family also includes herbs and climbers. Recent phylogenetic studies have lead to the inclusion in the Araliaceae of the herbaceous Hydrocotyloideae, previously included in the Apiaceae. Members of this group include Hemiphues, a genus of small montane herbs with creeping stems forming compact patches and more or less fleshy leaves found in New Zealand and Tasmania (Allan 1961). Didiscus is an Australian genus of erect, somewhat hairy herbs with flowers in an umbel. Neosciadium glochidiatum has leaves with the blade gradually constricted into a petiole and dense flower heads that start subglobular and elongate with maturity. Uldinia ceratocarpa has palmately divided leaves and minutely papillate or reticulate mericarps.

Other representatives of Araliaceae include the woody climbers in the genus Hedera, the best known of which is the garden ivy Hedera helix. Dendropanax is a genus of shrubs and trees with simple leaves found in eastern Asia and the Americas (Liogier 1995). Brassaia is an Indo-Pacific genus of few-branched trees with pinnately compound leaves and sessile flowers with seven to eighteen (usually twelve) locules (Liogier 1995). Oreopanax is a Neotropical genus of shrubs and trees with flowers borne in heads arranged into a raceme or panicle.

Ginseng and ivy
Published 24 October 2011

The Araliaceae are a family of nearly 1500 species of flowering plants found around the world, but primarily in the Old World tropics. Most of its members are trees or shrubs, but there are also some herbaceous or climbing species. Many Araliaceae have palmate leaves, and they often produce inflorescences in umbels. Not that many Araliaceae hold much economic prominence: Tetrapanax papyriferus is used to make rice paper, while the genus Panax includes the ginsengs that are widely regarded as something of a wonder-drug for no apparent good reason. Some other species are well known as garden plants, such as ivy Hedera helix. Back in my home country of New Zealand, Araliaceae include some of the most familiar small native trees such as pate Schefflera digitata and the five-fingers and lancewoods of the genus Pseudopanax.

A young lancewood Pseudopanax crassifolius, photographed by Mike Hudson. Lancewood is notable for its differing growth habits over its lifespan: this individual is just beginning to change from its juvenile to its mature foliage. When the plant is young, the long, narrow, tooth-edged leaves hang down around the trunk. As the tree matures, it produces leaves that are shorter, broader and with less strong teeth, and that are held upwards and outwards. The juvenile and mature trees are so different in appearance that they were initially described as different species.

The Araliaceae have long been recognised as close relatives of the Apiaceae, the family including carrots and celery, to the extent that some authors have combined the two in a single family. Most recent researchers have maintained the distinction, but phylogenetic studies have indicated that some genera previously treated within the Apiaceae, notably the water and marsh pennyworts of the genus Hydrocotyle, are better treated as basal Araliaceae (Plunkett et al. 1997). Relationships within the Araliaceae are somewhat less straightforward, as molecular phylogenetic studies have indicated that there has been a great deal of homoplasy in morphological characters (Plunkett et al. 2004). Some of the larger genera in the family (notably the genus Schefflera, to which nearly half the species of Araliaceae have been assigned) appear to be significantly polyphyletic, some of them not even resolving in particularly proximate clades. The difficult nature of many araliaceous genera has long been realised: in 1868, the botanist Berthold Seemann referred to the then-poorly defined Panax as “one of the great lumber rooms of our science” (Wen et al. 2001).

American ginseng Panax quinquefolius, from here. Red ginseng is derived from the root of this species and the Asian P. ginseng; however, over-harvesting has lead to the endangerment of wild populations of the latter.
Systematics of Araliaceae

Characters (from Kårehed 2003): Mostly woody (sometimes herbaceous); leaves usually alternate (sometimes opposite or whorled), simple or compound, penninerved or palmatinerved, serrate or entire, domatia absent; stipules absent or present; petiole with or without sheathing base; stomata para-, aniso- or anomocytic; hypodermis present or absent. Inflorescences mostly simple umbels; flowers usually with articulated pedicels, unisexual or hermaphroditic, pentamerous (sometimes 3-12-merous, stamens and carpels sometimes twice that number or up to ca. 200), petals free (sometimes connate at base); petals single-veined; aestivation imbricate or valvate; disc present; style mostly with 3-4 free stylodia, sometimes connate into style; stylopodium mostly present; ovary inferior, 2-5-locular (occasionally plurilocular); fruit a drupe or berry. Chromosome number x=11-12+. Secretory canals present; nodes trilacunar (rarely unilacunar); perforation plates simple and/or scalariform; vessels mostly solitary or mainly in groups, sometimes with helical thickenings, with simple pits or pits with very narrow borders; axial parenchyma scanty, paratracheal; rays heterogeneous; fibres with simple pits, mostly septate. Pollen colporate; exine usually reticulate.

<==Araliaceae
    |--Hydrocotyloideae [Hydrocotylaceae, Hydrocotyleae]HH00
    |    |--HydrocotyleK03
    |    |--TrachymeneT00
    |    |--DidiscusDK-DW00
    |    |    |--D. albiflorusM89
    |    |    |--D. hemicarpusC16
    |    |    `--D. pusillaDK-DW00
    |    |--Hemiphues Hooker 1847A61
    |    |    `--H. suffocata Hooker 1847A61 [=Actinotus suffocataA61, Hemiphues bellidioides var. suffocataC06]
    |    |         |--H. s. var. suffocataA61
    |    |         `--H. s. var. novae-zelandiae (Petrie) Allan 1961 (see below for synonymy)A61
    |    |--Neosciadium glochidiatumLP16, K90
    |    `--Uldinia ceratocarpaLP16, K90
    `--+--+--Fatsia japonicaDK-DW00
       |  |--KalopanaxDK-DW00
       |  |    |--K. pictusT03
       |  |    `--K. septemlobus [incl. Acanthopanax ricinifolia]LO98
       |  |--HederaK03
       |  |    |--H. canariensisRA07
       |  |    |--H. helixK03
       |  |    `--H. nepalensisD07
       |  |         |--H. n. var. nepalensisLP16
       |  |         `--H. n. var. sinensisLP16
       |  `--+--Dendropanax Decne. & Planch. 1854DK-DW00, L95
       |     |    |--D. arboreus (L.) Decne. & Planch. 1854 (see below for synonymy)L95
       |     |    |--D. laurifolius (March.) Decne. & Plant. ex Schneider 1909 (see below for synonymy)L95
       |     |    |--D. selleanusJ87
       |     |    `--D. trifidusLP16
       |     `--OreopanaxDK-DW00
       |          |--O. capitatumJ87
       |          |--O. floribundumF11
       |          |--O. peltatumBTA75
       |          `--O. sanderianusDK-DW00
       `--+--PanaxK03
          `--+--ScheffleraK03
             `--AraliaK03
Araliaceae incertae sedis:
  PseudopanaxA61
  TetraplasandraLP16
    |--T. gymnocarpaK03
    `--T. kavaiensisK03
  Heptapleurum venulosumV02, P03
  PolysciasP95
  OsmoxylonP95
  GastoniaP95
    |--G. serratifolia (Miq.) Philipson 1979 (see below for synonymy)P95
    `--G. spectabilis (Harms) Philipson 1970 (see below for synonymy)P95
  Tieghemopanax sambucifoliusYY22, C70
  HeteropanaxYY22
  BrassaiopsisYY22
    |--B. ciliateLP16
    |--B. glomerulataDD73
    |    |--B. g. var. glomerulataDD73
    |    `--B. g. var. serrataDD73
    |--B. hainlaDD73
    `--B. speciosaLP16
  GumbleaYY22
  PorospermumYY22
  StrobilopanaxYY22
  Acanthopanax cissifoliusO88
  Gilibertia trifidaMH98
  Brassaia Endl. in Endl. Fenzl. 1839L95
    `--B. actinophylla Endl. in Endl. & Fenzl. 1839 (see below for synonymy)L95
  OplopanaxNS06
  Anakasia simplicifolia Philipson 1973P95
  HarmsiopanaxP95
    |--H. harmsii Sch. ex Sch. & Laut. 1905P95
    `--H. ingens Philipson 1973P95
         |--H. i. ssp. ingensP95
         `--H. i. ssp. moniliformis Philipson 1973P95
  PlerandraP95
    |--P. brassii Philipson 1951P95
    |--P. solomonensis Philipson 1951P95
    `--P. stahliana Warb. 1894P95
  ArthrophyllumP95
    |--A. macranthum Philipson 1951P95
    |--A. pacificum Philipson 1977P95
    `--A. proliferum Philipson 1977P95
  Meryta Forster & Forster 1776A61
    |--M. colorata Bailey 1898P95
    `--M. sinclairii (Hooker) Seem. 1862 [=Botryodendrum sinclairii Hooker 1853]A61
  DidymopanaxJ87
    |--D. tremulumJ87
    `--D. vinosumMM96
  Raukaua Seemann 1866 [=Raukana]MFH97
    |--*R. edgerleyi (Hooker) Seemann 1866MFH97, A61 (see below for synonymy)
    |--R. anomalus (Hooker) Mitchell, Frodin & Heads 1997MFH97 (see below for synonymy)
    |--R. × parvus (Kirk) Heenan 1998 (see below for synonymy)H98
    |--R. × serratus (Kirk) Heenan 1998 (see below for synonymy)H98
    `--R. simplex (Forster) Mitchell, Frodin & Heads 1997MFH97 (see below for synonymy)
         |--R. s. var. simplexMFH97
         `--R. s. var. sinclairii (Hooker) Mitchell, Frodin & Heads 1997 (see below for synonymy)MFH97
  Kirkophytum (Harms) Allan 1961 [=Stilbocarpa sect. Kirkophytum Harms 1894]A61
    |--*K. robustum (Kirk) Allan 1961 (see below for synonymy)A61
    `--K. lyallii (Armstrong) Allan 1961 [=Stilbocarpa lyallii Armstrong 1881, Aralia lyalli Kirk 1885]A61
  Boerlagiodendron Harms 1894 (see below for synonymy)S62
    |--B. barbatumS62
    `--B. orientale Guillaumin 1938S62
  AstrotrichaB00
    |--A. floccosaM94
    |--A. ledifoliaB96
    |--A. longifoliaB00
    `--A. paucifloraB00
  Araliopsoides brevilova Berry 1916CBH93
  PentapanaxCBH93
  ‘Tricolporopollenites’ armatusCBH93
  CussoniaLP16
    |--C. angolensisLP16
    |--C. natalensisLP16
    |--C. paniculataDK-DW00
    `--C. spicataLP16
  CheirodendronMFH97
    |--C. bastardianum [=Nothopanax bastardianus; incl. C. marquesense]MFH97
    `--C. trigynumMFH97
  EleutherococcusMFH97
  MerrilliopanaxLP16
    |--M. chinensisLP16
    `--M. listeriDD73
  HederopsisT-W89
  ChengiopanaxLP16
  MotherwelliaLP16
  TetrapanaxLP16
  Trevesia palmataLP16
  MacropanaxLP16

Boerlagiodendron Harms 1894 [=Eschweileria Zippel ex Boerlage 1887 non Eschweilera Martius ex DC. 1828; incl. Unjala Reinw. ex Boerlage 1887]S62

Brassaia actinophylla Endl. in Endl. & Fenzl. 1839 [=Schefflera actinophylla (Endl.) Harms in Engl. & Prantl 1894]L95

Dendropanax arboreus (L.) Decne. & Planch. 1854 [=Aralia arborea L. 1759, Gilibertia arborea (L.) Marchal ex Durand & Pittier 1891, Hedera arborea (L.) Sw. 1797; incl. Sciadophyllum jacquini Griseb. 1860]L95

Dendropanax laurifolius (March.) Decne. & Plant. ex Schneider 1909 [=Gilibertia laurifolia March. in Urban 1899]L95

Gastonia serratifolia (Miq.) Philipson 1979 [=Arthrophyllum serratifolium Miq. 1861; incl. G. papuana Miq. 1863, Polyscias papuana (Miq.) Seem. 1865, Tetraplasandra paucidens Miq. 1863, T. solomonensis Philipson 1951]P95

Gastonia spectabilis (Harms) Philipson 1970 [=Peekeliopanax spectabilis Harms 1926; incl. G. boridiana Harms 1938]P95

Hemiphues suffocata var. novae-zelandiae (Petrie) Allan 1961 [=H. novae-zelandiae Petrie 1880, Actinotus novae-zelandiae Petrie 1881]A61

*Kirkophytum robustum (Kirk) Allan 1961 [=Aralia lyallii var. robusta Kirk 1891, Stilbocarpa robusta (Kirk) Cockayne 1909; incl. S. bollonsii Cockayne ex Watson 1909 (n. n.)]A61

Raukaua anomalus (Hooker) Mitchell, Frodin & Heads 1997MFH97 [=Panax anomalum Hooker 1843MFH97, Neopanax anomalum (Hooker) Allan 1961A61, Nothopanax anomalum (Hooker) Seemann 1866A61, Pseudopanax anomalum (Hooker) Koch 1859MFH97; incl. Panax microphyllum Colenso 1884MFH97, Nothopanax microphyllumC06, Pa. anomalum var. microphyllum (Colenso) Kirk 1899MFH97]

*Raukaua edgerleyi (Hooker) Seemann 1866MFH97, A61 [=Panax edgerleyi Hooker 1853A61, Nothopanax edgerleyi (Hooker) Harms 1894A61, Pseudopanax edgerleyi (Hooker) Koch 1859MFH97]

Raukaua × parvus (Kirk) Heenan 1998 [R. anomalus × R. simplex; =Panax simplex var. parvus Kirk 1899, Nothopanax parvum (Kirk) Cockayne 1909; incl. N. × anomosimplex Allan 1927 (n. n.), N. × simpan Allan 1926 (n. n.), N. × simpanomalum Cockayne 1929 (n. n.)]H98

Raukaua × serratus (Kirk) Heenan 1998 [R. edgerleyi × R. simplex; =Panax edgerleyi var. serratus Kirk 1889, R. edgerleyi var. serratus (Kirk) Mitchell, Frodin & Heads 1997]H98

Raukaua simplex (Forster) Mitchell, Frodin & Heads 1997MFH97 [=Panax simplex Forster 1786MFH97, Neopanax simplex (Forster) Allan 1961A61, Nothopanax simplex (Forster) Seemann 1866A61, Pseudopanax simplex (Forster) Koch 1859MFH97; incl. No. integrifoliumC06, Panax integrifolius Colenso 1888C06, Pa. simplex var. quercifolius Kirk 1899MFH97, Nothopanax simplex var. quercifolium Kirk 1899A61]

Raukaua simplex var. sinclairii (Hooker) Mitchell, Frodin & Heads 1997 [=Panax sinclairii Hooker 1864, Neopanax simplex var. sinclairii (Hooker) Allan 1961, Nothopanax sinclairii (Hooker) Seem. 1866, Pseudopanax simplex var. sinclairii (Hooker) Edgar 1973]MFH97

*Type species of generic name indicated

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