[Berberidopsidales+Caryophyllales]
Berberidopsidanae (Doweld) Thorne et Reveal in Bot. Rev. 73: 85. Apr-Jun 2007; Aextoxicales Takht. ex Reveal in Kew Bull. 66: 47. Mar 2011
Habit Bisexual, evergreen climbing shrubs or lianas, or dioecious evergreen tree.
Vegetative anatomy Phellogen ab initio superficial or absent. Vessel elements with scalariform perforation plates; lateral pits scalariform, opposite or alternate, simple pits. Imperforate tracheary xylem elements tracheids or fibre tracheids with bordered pits, non-septate (sometimes also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular; multiseriate parts with procumbent cells, uniseriate parts with quadrate to upright cells. Axial parenchyma apotracheal diffuse, or paratracheal scanty, or absent. Sieve tube plastids ? type. Nodes 3:3, trilacunar with three leaf traces. Sclereids sometimes present in all vegetative parts. Prismatic calciumoxalate crystals sometimes abundant; druses often present. Wood rays and axial parenchyma with dark-staining substances.
Trichomes Hairs unicellular or multicellular, uniseriate, sometimes peltate-lepidote.
Leaves Alternate (spiral), simple, entire, with involute or conduplicate (and sometimes slightly peltate) ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection annular. Venation pinnate or palmate. Stomata cyclocytic. Cuticular waxes? Rhomboidal calciumoxalate crystals and/or druses few to numerous. Leaf margin entire or spinose-serrate.
Inflorescence Terminal or axillary, raceme or spike, or flowers solitary terminal.
Flowers Actinomorphic. Hypogyny. Outer and inner tepals different or outer tepals sepaloid and inner ones petaloid. Tepals whorled with outer tepals (four or) five (or six), with imbricate aestivation, caducous, and inner tepals (four or) five (or six), with imbricate aestivation, persistent; or tepals spiral or whorled and 13 to 17, free.
Androecium Stamens usually five to numerous, in one or two whorls or spiral. Filaments thick, free from each other and from tepals. Anthers dorsifixed to subbasifixed, non-versatile, tetrasporangiate, introrse or latero-introrse, longicidal to almost poricidal (dehiscing by longitudinal slits or short subapical pore-like slits). Tapetum secretory? Female flowers with fleshy staminodia alternating with nectariferous glands.
Pollen grains Microsporogenesis simultaneous? Pollen grains tricolpate or tricolporate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, imperforate or microperforate, foveolate, regulate or striate.
Gynoecium Pistil composed of two, three or five connate carpels. Ovary superior, usually unilocular (sometimes bilocular with one locule usually sterile). Style single, simple. Stigma bilobate, type? Male flowers with pistillodium.
Ovules Placentation usually parietal (sometimes apical). Ovules two or more per carpel, pleurotropous or anatropous, apotropous or epitropous, bitegmic, crassinucellar. Micropyle bistomal or endostomal. Nucellar beak sometimes present. Megagametophyte monosporous, Polygonum type. Endosperm development cellular or nuclear? Endosperm haustoria? Embryogenesis?
Fruit A berry or a dry one-seeded drupe.
Seeds Aril absent. Seed coat testal (often endotestal). Exotesta often fleshy or leathery. Endotesta well developed. Tegmen often persistent. Perisperm not developed. Endosperm oily and proteinaceous (sometimes ruminate). Embryo usually small (sometimes long and curved), chlorophyll? Cotyledons two. Germination?
Cytology n = 16 (Aextoxicon)
DNA d copy of nuclear gene RPB2 lost.
Phytochemistry Cyanidin, tannins, proanthocyanidins, cyclopentenoid cyanogenic glycosides, and lotaustralin present. Cyanogenesis via isoleucine or valine. Tannins sometimes abundant.
Systematics Berberidopsidales are sister-group to Asteranae, according to Qiu & al. (2010) and Moore & al. (2011). Aextoxicon is sister to Berberidopsidaceae with high bootstrap support.
AEXTOXICACEAE Engl. et Gilg |
Genera/species 1/1
Distribution Southern and Central Chile and adjacent parts of Argentina.
Fossils Unknown.
Habit Dioecious, evergreen tree.
Vegetative anatomy Phellogen ab initio superficial. Medulla heterogenous. Vessel elements with scalariform perforation plates; lateral pits scalariform, simple pits. Imperforate tracheary xylem elements tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular; multiseriate parts with procumbent cells, uniseriate parts with quadrate to upright cells. Axial parenchyma apotracheal diffuse, or paratracheal scanty. Sieve tube plastids ? type. Nodes 3:3, trilacunar with three leaf traces. Sclereids present in all vegetative parts of the plant. Prismatic (rhomboidal) calciumoxalate? crystals abundant; druses scarce. Wood rays and axial parenchyma with dark-staining substances.
Trichomes Younger branches covered by ferrugineous peltate lepidote hairs.
Leaves Alternate (spiral) or almost opposite, simple, entire, with conduplicate (and sometimes slightly peltate) ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection annular. Petiole slightly pulvinate. Venation pinnate. Stomata cyclocytic to actinocytic, with five to seven subsidiary cells. Cuticular waxes? Lower side of leaf covered by ferrugineous peltate lepidote hairs. Sclereids of lamina approx. half as long as leaf thickness. Rhomboidal crystals and druses present. Leaf margin entire. Leaf primordia not enclosed by bud scales during winter.
Inflorescence Usually axillary, raceme with three or more flowers. Branches and flowers covered by ferrugineous peltate lepidote hairs. Floral prophylls (bracteoles) enclosing floral bud (calyptrate).
Flowers Actinomorphic. Bud covered by two congenitally fused calyptrate floral prophylls (bracteoles). Hypogyny. Outer tepals (‘sepals’) (four or) five (or six), decussate to spiral, with imbricate aestivation, thin, bract-like, caducous, free. Inner tepals (‘petals’) (four or) five (or six), mainly spiral, with imbricate aestivation, clawed, persistent, free. Nectariferous receptacular disc as (four or) five (or six) reniform glands, alternating with stamens.
Androecium Stamens (four or) five (or six), antesepalous. Filaments thick, free from each other and from petals. Anthers dorsifixed to subbasifixed, non-versatile, tetrasporangiate, introrse, intermediary between longicidal and poricidal (dehiscing by short subapical pore-like slits). Tapetum secretory? Female flowers with fleshy staminodia alternating with nectariferous glands.
Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, ?-cellular at dispersal. Exine?, with ? infratectum, sculpturing?
Gynoecium Pistil composed of usually one carpel (rarely two carpels). Ovary superior, unilocular. Style single, simple, short, curved at one side and pressed against ovary, with bifid apex. Stigmatic branches two, type? Male flowers with pistillodium.
Ovules Placentation apical-marginal. Ovules two per carpel, anatropous, pendulous, apotropous, bitegmic, crassinucellar. Micropyle endostomal, with very prolonged endostoma. Outer integument two or three cell layers thick, interrupted, hood-like. Inner integument five to seven cell layers thick. Obturator present. Megasporangium well developed, with strongly prolonged apex, nucellar beak, reaching outside integuments. Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustoria? Embryogenesis?
Fruit A dry one-seeded drupe with leathery pericarp. Endocarp usually dehiscing along two vertical lines.
Seeds Aril absent. Caruncle present. Seed coat testal. Testa tanniniferous, approximately six cell layers thick and with thin cell walls. Endotesta well developed. Tegmen weakly developed? Perisperm not developed. Endosperm ruminate, oily and proteinaceous. Embryo long, curved, well differentiated, transverse relative to longitudinal axis of seed, chlorophyll? Cotyledons two, flattened, cordate-orbicular. Germination?
Cytology n = 16
DNA
Phytochemistry Virtually unknown. Tannins abundant. Aluminium possibly accumulated?
Use Timber.
Systematics Aextoxicon (1; A. punctatum; from 36° to 44°S in Chile and adjacent parts of Argentina, the Chilean Coastal Cordillera).
Aextoxicon is sister to Berberidopsidaceae.
There is no clear delimitation in the perianth between the outer, sepaloid, tepals (sepals) and the inner, petaloid, tepals (petals).
BERBERIDOPSIDACEAE (Veldkamp) Takht. |
Genera/species 2/3
Distribution Central Chile, easternmost Australia.
Fossils Unknown.
Habit Bisexual, evergreen climbing shrubs or lianas.
Vegetative anatomy Phellogen absent. Vessel elements with scalariform perforation plates; lateral pits almost absent, opposite or alternate, simple pits. Imperforate tracheary xylem elements fibre tracheids with bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular; multiseriate parts with procumbent cells, uniseriate parts with quadrate to upright cells. Axial parenchyma very rare (apotracheal diffuse or paratracheal vasicentric) or absent. Sieve tube plastids S? type. Nodes 3:3, trilacunar with three leaf traces. Prismatic calciumoxalate crystals and druses abundant. Wood rays (and axial parenchyma) with dark-staining substances.
Trichomes Hairs in Berberidopsis unicellular to tricellular, uniseriate.
Leaves Alternate (spiral), simple, entire, with involute ptyxis (Berberidopsis). Stipules and leaf sheath absent. Petiole vascular bundle transection annular. Petiole with druses and single (rhomboidal?) crystals, sometimes pulvinate. Venation palmate. Stomata cyclocytic (sometimes bicyclic). Cuticular waxes? Leaf margin entire to spinose-serrate.
Inflorescence Terminal, spike or flowers solitary.
Flowers Actinomorphic. Hypogyny. Tepals 5+5 (Streptothamnus) or 12 to 14(–17) (Berberidopsis), free, whorled (Streptothamnus) or spiral (Berberidopsis, in groups of five tepals each), sepals and petals different (Streptothamnus) or outer tepals sepaloid and inner ones grading into petaloid tepals (Berberidopsis). Nectariferous disc extrastaminal, lobed (possibly absent in Streptothamnus), sometimes persistent.
Androecium Stamens six to numerous, either six to 13 in one whorl (Berberidopsis) or numerous more or less spiral (Streptothamnus). Filaments free from each other and from tepals. Anthers subbasifixed (in Berberidopsis adnate along lower part of connective; in Streptothamnus more or less basifixed), non-versatile, tetrasporangiate, latero-introrse, longicidal (dehiscing by longitudinal slits); connective somewhat prolonged. Tapetum secretory? Staminodia absent?
Pollen grains Microsporogenesis simultaneous? Pollen grains tricolpate or tricolporate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, imperforate or microperforate, foveolate to regulate, striate.
Gynoecium Pistil composed of three to five (rarely six, in Streptothamnus four) connate carpels. Ovary superior, unilocular. Style single, simple, thick, hollow. Stigma punctate, capitate or slightly lobate, type? Pistillodium absent.
Ovules Placentation parietal (with three or five placentae). Ovules two or more per carpel (ovary unilocular), anatropous, pleurotropous or epitropous, bitegmic, crassinucellar. Micropyle endostomal or bistomal. Outer integument approx. four cell layers thick. Inner integument approx. four cell layers thick. Megagametophyte monosporous, Polygonum type? Endosperm development nuclear? Endosperm haustoria? Embryogenesis?
Fruit A berry with persistent style and, in Streptothamnus, persistent calyx.
Seed Aril absent. Chalazal arilloid (caruncle?) present in Streptothamnus. Seed with prolonged raphe, in Streptothamnus with chalazal arilloid. Seed coat testal. Exotesta fleshy or leathery, with enlarged cells. Inner mesotestal cells often with sclereids. Endotestal cells with single crystals, palisade, with lignified walls. Exotegmen often weakly developed, fibrous, with lignified cell walls. Endotegmen fairly persistent. Perisperm not developed. Endosperm copious, oily and proteinaceous. Embryo small, chlorophyll? Cotyledons two. Germination?
Cytology n = 21?
DNA
Phytochemistry Cyanidin, proanthocyanidin, lotaustralin, and cyclopentenoid cyanogenic glycosides present. Cyanogenesis via isoleucine or valine. Ellagic acid not found.
Use Ornamental plants (Berberidopsis corallina).
Systematics Berberidopsis (2; B. corallina: central Chile; B. beckleri: southeastern Queensland, northeastern New South Wales), Streptothamnus (1; S. moorei; southeastern Queensland, northeastern New South Wales).
Berberidopsidaceae are sister-group to Aextoxicon (Aextoxicaceae). The phylogenetic position of Streptothamnus is disputed (see, e.g., Ronse DeCraene 2017), and molecular analyses are needed. Streptothamnus moorei may even be more distantly allied to Berberidopsis.
Literature
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