VITACEAE (vinrankaväxter) 12-17/750-800 VITALES

Takhtajan, Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 264. 4 Feb 1967

VITACEAE Juss.

Vitales Juss. ex Berchtold et J. Presl, Přir. Rostlin: 225. Jan-Apr 1820 [‘Vites’]; Leeaceae (DC.) Dumort., Anal. Fam. Plant.: 21, 27. 1829, nom. cons.; Ampelopsidaceae Kostel., Allg. Med.-Pharm. Fl.: 1194. Jan-Oct 1835 [’Ampelopsideae’]; Leeales DC. in C. F. P. von Martius, Consp. Regn. Veg.: 26. Sep-Oct 1835 [‘Leeaceae’]; Cissaceae Drejer, Comp. Medic. Bot.: 45. 1840 [’Cisseae’]; Pterisanthaceae J. Agardh, Theoria Syst. Plant.: 268. Apr-Sep 1858 [’Pterisantheae’]; Vitanae Takht. ex Reveal in Novon 2: 237. 13 Oct 1992

Distribution Mainly in tropical and subtropical regions, some species in warm-temperate areas; Leea: Southeast Asia, Malesia, New Guinea and Australia, two species in Africa and Madagascar.

Fossils Fruits assigned to Vitaceae are known from c 66 My old sediments in India (Manchester & al. 2013). Seeds are abundant in Cenozoic layers from the Paleocene onwards, and the clade was well differentiated during the early Eocene. Palaeogene fossils have been assigned to several extant genera.

Habit Bisexual, monoecious, polygamomonoecious, dioecious or polygamodioecious, usually evergreen (sometimes deciduous) lianas (sometimes climbing, perennial herbs) with leaf-opposed, simple or branched branch-tendrils with small adhesive cushions attaching plant to trees or cliffs etc. (rarely small succulent trees with swollen stem; Leea consists of evergreen trees, shrubs or perennial herbs without tendrils). Lenticels often abundant and significant.

Vegetative anatomy CAM physiology sometimes occurring? Phellogen ab initio superficially or deeply seated. Medulla continuous or interrupted by diaphragms at nodes. Primary medullary strands wide. Secondary lateral growth normal or anomalous (in Tetrastigma from concentric cambia). Cambium often stratified. Vessel elements usually with simple (rarely scalariform) perforation plates; lateral pits scalariform, opposite or alternate, simple pits. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, usually septate. Wood rays multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty vasicentric, or banded, or absent. Wood partially storied. Tension wood absent. Secondary phloem often stratified into hard fibrous and soft parenchymatic Tyloses present. zones. Sieve tube plastids Pcs type (P2b type), with several polygonal protein crystals and starch. Nodes 3–7:3–7, trilacunar to multilacunar with three to seven leaf traces, often swollen. Calciumoxalate as druses, raphides, or single prismatic crystals (raphides in Leea with filiform appendages). Parenchyma with raphide sacs each containing often hundreds of raphides. Shoot apex producing foliar primordia and primordia developing into inflorescences or tendrils.

Trichomes Hairs unicellular or multicellular, uniseriate or branched, sometimes furcate, often lepidote, peltate, and vesicular hairs; sugar- and oil-storing food bodies, pearl glands (extrafloral nectaries; at least in Leea, Cissus and Vitis): multicellular, spherical, with short multiseriate stalk, caducous, sometimes with apical stoma.

Leaves Usually alternate (usually distichous; in Leea usually spiral, rarely opposite), simple or palmately compound, entire or palmately lobed (in Leea simply or repeatedly compound), with conduplicate ptyxis. Stipules usually pairwise, intrapetiolar, caducous or persistent (sometimes adaxially connate; rarely absent; in Leea sheathing, as wing-like outgrowths along petiole margin); leaf sheath absent. Petiole vascular bundle transection annular. Venation usually palmate (rarely pinnate). Stomata anomocytic, staurocytic, hemiparacytic or cyclocytic (Leea, or actinocytic?). Cuticular wax crystalloids usually ? (sometimes as tubular rodlets). Domatia as pockets or hair tufts, or absent. Mesophyll with numerous tanniniferous idioblasts, and raphide cells (raphide sacs) usually with both mucilage and calciumoxalate raphides, sometimes also mucilaginous idioblasts without raphides. Lamina often gland-dotted (with caducous pearl glands) and with lepidote and peltate hairs. Leaf margin serrate; teeth often coarse; leaf tooth with distally expanding gland and with foramen, and with lateral veins running from above and below tooth (in Leea with small glandular apex and one lateral vein running further above tooth).

Inflorescence Usually leaf-opposed (sometimes terminal or axillary; in Leea usually terminal), cymose corymb or panicle (rarely spike-like; in Leea often umbel-like). Flowers often replaced by tendrils (sometimes with specialized ‘sucking discs’). Inflorescence in Leea often densely beset with ferrugineous hairs.

Flowers Actinomorphic, small. Hypanthium present or absent (absent in, e.g., Leea). Usually hypogyny (in Leea rarely half epigyny). Sepals (three or) four or five (to seven), usually with open (in Leea valvate) aestivation, often as teeth or scales or reduced to ring, caducous (or sometimes persistent?), connate. Petals (three or) four or five (to seven), with valvate aestivation, usually free (in Leea connate at base), in Vitis calyptrate (at apex connate forming hood, calyptra, detached at anthesis). Gynoecial nectaries present. Nectariferous disc intrastaminal, usually annular (nectary sometimes enclosing ovary, or as four or five nectariferous glands alternating with stamens, or absent; in Leea nectariferous disc absent, nectar not produced).

Androecium Stamens (three or) four or five (to seven), obhaplostemonous, alternisepalous, antepetalous; stamens and petals developed from common primordium. Filaments usually free from each other and from tepals (in Leea basally adnate to corolla tube [epipetalous] and upwards connate into anther tube, sometimes together with entire or bifid nectary lobes alternating with anthers, sometimes with pendant tubular membrane near centre of anther tube). Anthers dorsifixed, sometimes versatile, usually tetrasporangiate (rarely disporangiate), introrse, longicidal (dehiscing by longitudinal slits). Tapetum usually secretory (sometimes amoeboid-periplasmodial). Female flowers sometimes with staminodia?

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (in Leea also triporate), shed as monads, usually bicellular at dispersal (in Leea usually tricellular). Exine semitectate, with columellate infratectum, usually reticulate (in Leea psilate).

Gynoecium Pistil composed of usually two (in Leea two or three [or four]) connate carpels, vertically or transversely orientated (odd carpel in Leea abaxial). Ovary usually superior (in Leea rarely semi-inferior), bilocular or trilocular (in Leea sometimes quadrilocular, each locule divided by secondary septum). Style single, usually very short (rarely absent; in Leea longer). Stigma usually capitate (in Tetrastigma quadrilobate), usually non-papillate (in Leea papillate), Dry type. Male flowers sometimes with pistillodium?

Ovules Placentation axile (seemingly subbasal-marginal; in Leea basal?). Ovules usually two per carpel (in Leea one per incompletely closed locule), anatropous, ascending, apotropous, bitegmic, crassinucellar. Micropyle usually endostomal (sometimes bistomal; in Leea exostomal). Outer integument usually four to seven cell layers thick. Inner integument usually two or three cell layers thick. Obturator placental or absent. Parietal tissue up to 17 cell layers thick. Nucellar cap present. Megagametophyte monosporous, Polygonum type. Egg apparatus in Cissus present outside? ovule. Endosperm development ab initio nuclear. Endosperm haustoria present or absent. Embryogenesis asterad.

Seeds Aril absent. Seed coat endotestal. Seeds perichalazal, with prominent adaxial raphe (flanked by two deep furrows) and abaxial chalazal tubercle. Testa with deep furrow adjacent to raphe. Vascular bundle more or less enclosing seed. Testa sometimes multiplicative. Cells with bundles of calciumoxalate raphides. Exotesta fleshy (sarcotesta). Mesotesta two to 17 cell layers thick. Endotesta two to five cell layers thick, lignified, usually with calciumoxalate raphides (rarely palisade). Exotegmen crossed tracheidal. Endotegmen mucilaginous. Perisperm not developed? Endosperm copious, oily and proteinaceous, sometimes ruminate, usually Y-, T-, M- or U-shaped in cross-section. Embryo minute, straight, well differentiated, without chlorophyll. Cotyledons two. Germination phanerocotylar.

DNA d copy of nuclear gene RPB2 lost. Mitochondrial genome in Vitis very large, with expanded intergenic spacers. Mitochondrial intron coxII.i3?

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), cyanidin, oleanolic acid derivatives, ellagic and gallic acid, hydrolyzable and non-hydrolyzable tannins, proanthocyanidins (prodelphinidins), caffeic acid, chlorogenic acid, alkaloids, naphthoquinones, tartric acid, and myo-inisitol present. Saponins and cyanogenic compounds not found.

Use Ornamental plants, fruits (grapes, raisins, juice, etc.), wines and liquors (Vitis), medicinal plants.

Systematics Vitaceae are sister-group to the remaining Rosidae with high support, according to multigene analyses including 17 genes (Soltis & al. 2011). Vitaceae were recovered as sister to [Saxifragales+Rosanae], although with weak support, in analyses of plastid inverted repeats in 244 taxa (Moore & al. 2011). On the other hand, Vitaceae were found to be sister-group to Saxifragales with relatively high support in analyses by Zhang & al. (2016).

Leeoideae Burmeist., Handb. Naturgesch.: 338. 1837 [‘Leeaceae’]

1/34–>70. Leea (34–>70; Southeast Asia, Malesia to New Guinea, tropical Australia, one species, L. guineensis, in Africa and Madagascar). – Evergreen trees, shrubs or perennial herbs. Raphides with filiform appendages. Leaves spirally arranged, simply or twice compound. Stipules sheathing, visible as wing-like outgrowths along petiole margin. Leaf teeth with small glandular apex and one lateral vein running further above tooth. Petals connate at base. Nectariferous disc absent, nectar not produced. Filaments adnate at base to corolla tube, upwards connate and forming tube together with “nectary” lobes. Ovary locules divided by secondary septum. Micropyle exostomal. Fruit a four- to six-seeded berry. Testa rarely with raphides. n = (10–)12, 24. –Leea is sister to the remaining Vitaceae.

Vitoideae Eaton, Bot. Dict., ed. 4: 40. Apr-Mai 1836 [‘Vites’]

c 12/690–890. 'Ampelopsis' (c 16; temperate and subtropical regions of Asia and America; paraphyletic), Nekemias (9; India, East and Southeast Asia, Java, Sulawesi, eastern North America), Cyphostemma (c 150; tropical and subtropical regions on both hemispheres, with their highest diversity in tropical Africa), 'Cayratia' (50–63; tropical regions in the Old World east to eastern Queensland; paraphyletic), Acareosperma (1; A. spireanum; Laos)?, Tetrastigma (90–95; tropical Asia to tropical Australia), ’Cissus’ (200–350; tropical and subtropical regions on both hemispheres; polyphyletic), Pterocissus (1; P. mirabilis; Hispaniola; in Cissus?), Clematicissus (2; C. angustissima: western Western Australia; C. opaca: eastern Queensland, eastern New South Wales), ‘Parthenocissus’ (10–18; temperate Asia, North America, Mexico; non-monophyletic), 'Ampelocissus' (95–120; tropical regions on both hemispheres; paraphyletic), Vitis (c 65; temperate and subtropical regions on the Northern Hemisphere). – Temperate to tropical regions on both hemispheres, with their largest diversity in tropical regions. Usually lianas (rarely trees or perennial herbs, sometimes with a swollen root), climbing with the aid of leaf-opposite tendrils evolved from inflorescence branches. Raphides glabrous. Leaves alternate (often distichous) or opposite. Cuticular wax crystalloids sometimes as tubular rods. Leaf teeth with gland distally expanded and with foramen; lateral veins running above and below tooth. Inflorescence usually leaf-opposite (rarely terminal). Hypogyny. Disc usually annular or as five glands (rarely surrounding gynoecium or absent). Pistil composed of two connate carpels, transversal or vertical. Style usually short (rarely long). Micropyle usually endostomal (sometimes bistomal). Outer integument four to seven cell layers thick. Inner integument (one or) two or three (or four) cell layers thick. Nucellar cap up to eight cell layers thick. Testa usually with raphides, sometimes multiplicative. n = 10–16, 19, 20. – In the plastid DNA sequence analysis by Ren & al. (2011), three distinct monophyletic groups were identified: 1) a “pentamerous clade” including Ampelocissus, Vitis and Parthenocissus; 2) a “tetramerous clade” containing Cissus, Cayratia and Tetrastigma; and 3) a clade sister to the two previous clades comprising the Ampelopsis lineage (including Rhoicissus and several species of Cissus, Lu & al. 2018). Clematicissus (not included in the analyses by Ren & al. 2011) is sister to Cissus striata and C. tweediana (Rossetto & al. 2007), and seems to be closely allied to the Ampelopsis lineage (Wen & al. 2007). Lu & al. (2018) discussed the Late Cretaceous rapid radiation of Vitoideae. Three main lineages were identified (supported by gynoecium morphology): 1) the Ampelopsis clade as sister to the rest, 2) Parthenocissus-Yua + Ampelocissus-Vitis, and 3) Cissus.

Phylogeny (simplified) of Vitaceae based on DNA sequence data (Ren & al. 2011)

Phylogeny (simplified) of Vitaceae based on DNA sequence data (Trias-Blasi & al. 2012)

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ROSIDAE Takht.

VITACEAE Juss.