[Sabiaceae+Proteales+[Trochodendrales+[Didymelales+Gunneridae]]]
Millingtoniaceae Wight et Arn., Prodr. Fl. Ind. Orient. 1: 115. 22 Sep 1834, nom. illeg.; Wellingtoniaceae Meisn., Plant. Vasc. Gen. 1: Comm.: 207, 346. 25-31 Oct 1840; Meliosmaceae (Endl.) Meisn., Plant. Vasc. Gen.: Comm.: 346. 2-4 Nov 1843 [‘Meliosmeae’], nom. rejic.; Sabiales Takht., in Divers. Classif. Fl. Pl.: 304. 24 Apr 1997; Sabianae Doweld, Tent. Syst. Plant. Vasc.: xxvii. 23 Dec 2001; Meliosmales C. Y. Wu in Acta Phytotaxon. Sin. 40: 312. 2002
Genera/species 3/c 90 or more than 140
Distribution The Himalayas, southern, eastern and southeastern Asia, Malesia to New Guinea, New Britain and Solomon Islands, tropical America north to central Mexico.
Fossils Sabia and Meliosma are represented by fossilized endocarps from the Turonian to Santonian and the Maastrichtian of Central Europe. Insitiocarpus includes endocarps from Cenomanian and Maastrichtian sediments in Central Europe and Late Turonian to Santonian wood from Japan has been assigned to Sabiocaulis sakuraii. Cenozoic records of leaves and endocarps are known from North America and Europe.
Habit Bisexual or polygamodioecious, usually evergreen (two species of Meliosma deciduous) trees, shrubs or lianas (in Sabia japonica with short spines).
Vegetative anatomy Phellogen ab initio superficial. Primary medullary strands wide and/or narrow. Vessel elements with simple and/or scalariform (sometimes reticulate) perforation plates; lateral pits alternate, simple or bordered pits. Imperforate tracheary xylem elements in Sabia tracheids and in Meliosma and Ophiocaryon fibre tracheids or libriform fibres with simple and/or bordered pits, septate or non-septate. Wood rays uniseriate or multiseriate, heterocellular or almost homocellular (absent in Sabia). Axial parenchyma apotracheal diffuse, paratracheal scanty vasicentric, or banded (rarely absent). Sieve tube plastids usually S type (sometimes also with protein crystals). Nodes 1:1, unilacunar with one leaf trace (Meliosma), complex. Secretory cavities absent. Wood rays sometimes with silica bodies or calciumoxalate. Prismatic crystals sometimes frequent.
Trichomes Hairs multicellular, usually simple (sometimes with bicellular apex).
Leaves Alternate (usually spiral, sometimes distichous), imparipinnate or seemingly simple (in reality unifoliolate), entire, often coriaceous, with conduplicate ptyxis (Meliosma). Foliar buds often perulate. Stipules and leaf sheath absent. Petiole vascular bundle transaction? Venation pinnate, brochidodromous. Stomata anomocytic or paracytic. Cuticular wax crystalloids absent. Lamina often with indistinct glandular dots. Mesophyll without secretory cavities. Calciumoxalate as abundant druses. Leaf margin serrate (sometimes spinose-serrate) or entire.
Inflorescence Terminal or axillary, compound, usually panicle (rarely raceme-like; flowers rarely solitary).
Flowers Actinomorphic or obliquely zygomorphic (Meliosma), small. Floral parts spiral (arranged 2:5). Hypogyny. Sepals (three to) five, with imbricate or open aestivation, free or connate at base (two inner sepals often smaller). Petals (three to) five, with usually imbricate (sometimes subvalvate) aestivation, antesepalous or alternisepalous, free (one or two inner petals in Meliosma and Ophiocaryon significantly smaller). Nectariferous disc intrastaminal, thin, annular, entire, dentate or lobate (often with glands), enclosing ovary base.
Androecium Stamens (four or) five (or six; Sabia) or two inner stamens fertile, with two basal outgrowths, and two or three outer staminodia (Meliosma, Ophiocaryon), antepetalous. Filaments often flattened, free, adnate at base to petals, often swollen or with annular extension (of connective?). Anthers dorsifixed, often versatile, disporangiate (thecae unilocellate), introrse or (seemingly) extrorse, longicidal (dehiscing by longitudinal lateral or transverse apical slits or in Ophiocaryon valves); connective in Meliosma and Ophiocaryon expanded. Anthers in Meliosma held under tension in bud by staminodia and dehiscing explosively. Tapetum secretory. Staminodia present (Meliosma, incurved and covering ovary and disk, often slightly connate at apex, and Ophiocaryon) or absent (Sabia). Secondary pollen display present; pollen grains caught on connective of own anther.
Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate to tricolporoidate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate or microreticulate.
Gynoecium Pistil composed of usually two (sometimes three) entirely or partially connate carpels (in Ophiocaryon apically free), often oblique relative to floral symmetry plane; carpel ascidiate, closed. Ovary superior, usually bilocular (rarely trilocular). Stylodia more or less connate into a style (stylodia in Ophiocaryon marginal), sometimes with a secretory canal. Stigmas usually two (sometimes three), capitate or punctate, non-papillate, Dry? or Wet type. Pistillodium usually absent.
Ovules Placentation usually axile (sometimes apical). Ovules usually two (sometimes one) per carpel, orthotropous or campylotropous, pendulous or horizontal, apotropous, unitegmic, crassinucellar. Micropyle absent, integument not covering megasporangium (Meliosma). Integument six cell layers thick (Sabia). Parietal cell developed from archesporial cell. Megagametophyte monosporous, Polygonum type. Endosperm development helobial. Endosperm haustorium chalazal (Sabia). Embryogenesis?
Fruit A drupe (Meliosma, Ophiocaryon) or schizocarp with one or two drupelets (Sabia), curved and flattened, with two drupaceous (Sabia) or nutlike one-seeded mericarps with persistent (adaxially subbasal) stylodia (one mericarp sometimes reduced). Fruit surface often strongly veined.
Seeds Aril absent. Seed with placental intrusion, condyle. Testa? Perisperm not developed. Endosperm small or absent. Embryo elongate, curved to coiled, oily (cf. the name ’snake nut’ of Ophiocaryon paradoxum), chlorophyll? Cotyledons two, usually folded (sometimes spirally twisted), oily. Hypocotyl large, curved to coiled (Ophiocaryon) or folded (Meliosma). Germination phanerocotylar.
Cytology n = 12, 16
DNA Nuclear gene paleoAP3 present.
Phytochemistry Flavonols (kaempferol, quercetin), pentacyclic triterpenoids, tannins, proanthocyanidins, and oleanolic acid derivatives present. Cyanogenic compounds not found.
Use Ornamental plants (Meliosma), vegetables, medicinal plants, timber, carpentries.
Systematics Meliosma (50 to more than 100; tropical Asia to Japan, Taiwan and New Guinea, Mexico, Central America, the West Indies, tropical South America), Ophiocaryon (10; tropical South America), Sabia (c 30; Southeast Asia, Malesia).
Sabia is sister to [Meliosma+Ophiocaryon] (Zúñiga 2015). There is low to relatively high bootstrap support for Sabiaceae being sister-group to Proteales (see, e.g., Moore & al. 2011; Soltis & al. 2011; Barniske & al. 2012), whereas other analyses identify Sabiaceae as sister to all Tricolpatae except Ranunculales and Proteales, or to Tricolpatae minus Ranunculales (Worberg & al. 2007). However, other positions have sometimes been found. Sabiaceae are sister to either Proteales or to the remaining Tricolpatae except Ranunculales. Awaiting more convincing results I treat them here as part of a basal Tricolpatae trichotomy.
Literature
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