CARYOPHYLLANAE Takht

Dilleniales DC. ex Bercht. et J. Presl, Přir. Rostlin: 223. Jan-Apr 1820 [‘Dilleniaceae’]; Soramiaceae Martinov, Tekhno-Bot. Slovar: 586. 3 Aug 1820 [‘Soramieae’]; Delimaceae Mart., Consp. Regni Veg.: 39. Sep-Oct 1835, nom. illeg.; Hibbertiaceae J. Agardh, Theoria Syst. Plant.: 200. Apr-Sep 1858; Dilleniidae Takht. ex Reveal et Takht. in Phytologia 74: 171. 25 Mar 1993, pro parte; Dillenianae Doweld, Tent. Syst. Plant. Vasc.: xxviii. 23 Dec 2001

Distribution Almost pantropical, with their largest diversity in tropical Asia and Australia; Hibbertia also in warm-temperate parts of Australia including Tasmania.

Fossils Fossilized seeds similar to Hibbertia and Tetracera are known from the Early Eocene of England and of Dillenia and Tetracera from the Late Eocene of Oregon (Chaney & Sanborn 1933). However, no unambiguous fossils of Dilleniaceae are known (Manchester & al. 2015).

Habit Usually bisexual (rarely dioecious), usually evergreen (rarely deciduous) trees or shrubs (rarely suffrutices; Tetracera consists of lianas; ‘Acrotrema’ costatum [nested in Dillenia] is a perennial herb with woody rhizome). Bark often intensely brown.

Vegetative anatomy Phellogen ab initio usually deeply (in, e.g., Dillenia superficially) seated. Primary stem with continuous vascular cylinder. Medulla sometimes septated by diaphragms. Concentric (successive) cambia, producing interxylary cylinders of secondary phloem, present in, e.g., Doliocarpus (causing anomalous secondary lateral growth). Vessel elements dimorphic, with usually scalariform or simple (sometimes opposite or reticulate) perforation plates (in Tetracera reticulate-scalariform); lateral pits scalariform, alternate or opposite, simple and/or bordered pits. Imperforate tracheary xylem elements tracheids with bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma usually apotracheal diffuse or diffuse-in-aggregates (sometimes paratracheal scanty, incompletely vasicentric or banded). Tension wood absent. Intraxylary phloem (concentric) present in some Dilleniaceae (e.g. Doliocarpus). Sieve tube plastids S type. Nodes usually ≥3:≥3, multilacunar with three or more leaf traces (up to 27:27; in Hibbertia 1:1, unilacunar with one trace, or 3:3, trilacunar with three traces). Heartwood sometimes with resin-like substances. Medulla often with brachysclereids or sclereids with white inclusions. Epidermal cell walls and wood often with silica bodies. Parenchymatic idioblasts with calciumoxalate raphides.

Trichomes Hairs unicellular or multicellular, uniseriate or branched, stellate, peltate, lepidote, often sclerified and/or silicified; glandular hairs very rare.

Leaves Usually alternate (spiral; very rarely opposite), simple (rarely pinnatifid), entire (rarely pinnatisect), often coriaceous, often harsh (in some species of Hibbertia ericoid; in some species of Pachynema scale-like on phyllocladia), with conduplicate ptyxis (leaves tending to elongate already as rolled-up). Stipules and leaf sheath absent. Petiole often with wing-like (sometimes stipule-like) appendages and, in some species of Hibbertia, perfoliate. Petiole vascular bundles? Venation craspedodromous, semicraspedodromous, brochidodromous or eucamptodromous; secondary veins often parallel, often reaching into leaf teeth; tertiary venation often scalariform. Stomata usually anomocytic (rarely paracytic, anisocytic or cyclocytic). Cuticular waxes absent? Domatia as pockets or hair tufts in some species. Epidermal cell walls often with silica crystals and sometimes also sclerified. Mesophyll often with sclerenchymatous idioblasts containing calciumoxalate as raphides, crystal sand or cuboidal to prismatic crystals. Leaf margin serrate or entire; leaf teeth with transparent glandular elongate apex.

Inflorescences Terminal or axillary, usually thyrsoid, panicle or botryoid, or flowers solitary. Floral prophylls (bracteoles) usually two (rarely one).

Flowers Usually actinomorphic (androecium in Didesmandra, Schumacheria and some species of Hibbertia horizontally zygomorphic), often large. Pedicel articulated at apex, persistent after anthesis. Usually hypogyny (rarely half epigyny). Sepals (three or) four or five (to 20), with imbricate aestivation (when five then quincuncial), spiral, persistent and often accrescent, free. Petals (two or) three to five (to seven), with imbricate aestivation (when five then quincuncial), spiral, often crumpled in bud, free, usually early caducous. Nectary absent. Disc absent.

Androecium Stamens (one or three to) five to more than 400 (in Hibbertia one to many more than 150, in Tetracera c. 50 to more than 500, in Dillenia ovalifolia to more than 900), often asymmetrical (in Pachynema flat and wide), spiral, reflexed in bud, often centrifugally arising from few stem bundles. Filaments free, or completely or only at base connate into one to five antesepalous fascicles (rarely forming a short tube), free from tepals. Anthers usually basifixed, non-versatile?, tetrasporangiate, introrse or latrorse, longicidal (dehiscing by longitudinal slits) or poricidal (dehiscing by apical pores or short slits); connective often enlarged and prolonged. Tapetum probably secretory (in some species possibly amoeboid-periplasmodial). Staminodia present in some species.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate (sometimes dicolpate or spiraperturate?; in Didesmandra and Schumacheria usually tetracolpate), tricolporate or triporate, shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate, reticulate or punctate.

Gynoecium Carpels one to ten (to 20), sometimes not completely closed, often antepetalous or median carpel adaxial, usually free (rarely connate at base; in Curatella half and in ’Neowormia’ completely connate), in usually one whorl (very rarely two whorls); compitum usually absent. Ovary usually superior (rarely semi-inferior), unilocular (apocarpy), or (bilocular to) quinquelocular to septalocular (syncarpy). Style (stylodia) one to ten (to 20), free. Stigmas usually capitate to punctate (in, e.g., Doliocarpus and Davilla peltate), non-papillate, Wet type. Pistillodia absent.

Ovules Placentation submarginal to basal (when one or two ovules and apocarpy) or axile to basal (when syncarpy). Ovules one to numerous (up to c. 20) per carpel, anatropous, campylotropous or amphitropous, when one ovule then apotropous (abaxially curved), when two ovules then one apotropous and one epitropous (adaxially curved), when at least four ovules then pleurotropous (laterally placed) and syntropous (curved with carpellary margin), bitegmic, crassinucellar. Micropyle exostomal or bistomal, Z-shaped (zig-zag). Outer integument two or three cell layers thick. Inner integument two to six cell layers thick. Parietal tissue six to 14 cell layers thick. Nucellar cap approx. two cell layers thick. Megasporocytes sometimes several. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis usually onagrad.

Fruit Usually a follicle or an assemblage of follicles (rarely a fleshy capsule, a berry or an assemblage of nutlets; fruit in Dillenia enclosed by fleshy enlarged calyx); filaments persistent.

Seeds: Oily or waxy and often lobate funicular aril usually present. Seed coat testal-tegmic. Testa non-multiplicative. Exotesta often fleshy (exotestal cells in Dillenia large, tanniniferous and gradually flattened). Endotesta palisade, usually lignified. Exotegmen usually tracheidal, usually with spiral or annular thickenings. Endotegmen tanniniferous. Cells with calciumoxalate raphide bundles often frequent. Perisperm not developed? Endosperm copious, oily, in Hibbertia also with starch grains. Zygote enveloped by membrane during early stages at least in Dillenia and Hibbertia. Embryo small, straight, well differentiated, without chlorophyll. Cotyledons two. Germination phanerocotylar.

Phytochemistry Flavonols (myricetin), flavonol sulphates, cyanidin, ellagic acid, gallic acid, hydrolyzable tannins, ellagitannins, proanthocyanidins (prodelphinidins), betulinic acid (mainly in bark), alkaloids (benzylisoquinoline alkaloids absent), and anthraquinones present. Cyanogenic compounds and saponins not found.

Systematics Dilleniaceae are identified as sister to Caryophyllales (Bell & al. 2010) or to the clade [Santalales+[Asteridae+[Berberidopsidales+Caryophyllales]]], i.e. to Superasteridae (Soltis & al. 2011, etc.). However, sequencing of the plastid inverted repeat for 244 species (Moore & al. 2011) recovered Dilleniaceae as sister to the clade [Superasteridae+Superrosidae], i.e. sister to the remaining Pentapetalae, with moderately high bootstrap support. Sun & al. (2016), in their whole plastid genome analysis of angiosperms, recovered Dillenia as sister to the rosids.

The absence of gynoecial compitum in numerous Dilleniaceae is obviously a secondary loss.

The subdivision below follows Horn (2009).

Delimeae DC., Syst. Nat. 1: 396, 397. 1-15 Nov 1817 [‘Delimaceae’]

1/c 50. Tetracera (c 50; pantropical). – Lianas. Vessel perforation plates scalariform or simple. Tertiary venation strongly percurrent, with well developed areoles. Stomata paracytic. Flowers actinomorphic. Stamens c. 50 to more than 500. Carpels isomerous relative to corolla. – Tetracera is sister to the remaining Dilleniaceae.

[Doliocarpeae+[Hibbertieae+Dillenieae]]

Doliocarpeae (Doliocarpoideae J. W. Horn in Intern. J. Plant Sci. 170: 809. 2009)

4–5/75–85. Davilla (18; tropical South America), Doliocarpus (50–60; southern Mexico, Central America, the West Indies, tropical South America), Curatella (1; C. americana; tropical America), Pinzona (1; P. coriacea; tropical America), Neodillenia (3; N. coussapoana, N. peruviana, N. venezuelana; Amazonia)? – Tropical America. Vessel perforation plates scalariform or simple. Tertiary venation strongly percurrent, with well developed areoles. Flowers actinomorphic. Stamens less than 200. Carpels relatively few. Internal compitum usually absent (present in Curatella and Pinzona). Stigma infundibular. Ovules two per carpel, collateral, one apotropous and one epitropous. – Doliocarpeae are sister-group to [Hibbertieae+Dillenioideae]. The position of Neodillenia is uncertain.

[Hibbertieae+Dillenieae]

Vessel perforation plates scalariform.

Hibbertieae Spach, Hist. Nat. Vég. 7: 413, 419. 4 Mai 1838

1/170–180. Hibbertia (170–180; Madagascar, Malesia to New Guinea, Australia, New Caledonia, Fiji, with their highest diversity in Australia). – Nodes 1:1, 3:3. Hairs sometimes stellate. Tertiary venation random; areoles poorly developed. Leaf margin usually serrate. Flowers usually actinomorphic (sometimes with androecial zygomorphy). Stamens one to c. 200 (rarely more), sometimes obdiplostemonous. Filaments usually free (outer ones sometimes connate at base). Outer stamens sometimes staminodial. Carpels relatively few. – Hibbertia is sister to Dillenieae.

Dillenieae DC., Syst. Nat. 1: 397, 411. 1-15 Nov 1817 [‘Dilleneae’]

3/c 105? Schumacheria (4; S. alnifolia, S. angustifolia, S. castaneifolia, S. raphanoides; Sri Lanka), Didesmandra (1; D. aspera; Borneo), Dillenia (c 100?; Indian Ocean islands, India, Sri Lanka, Southeast Asia, Malesia to New Guinea and tropical Australia). – Tropical Asia to tropical Australia. Phellogen superficial. Leaf base surrounding stem. Tertiary venation strongly percurrent, with well developed areoles. Flowers usually actinomorphic (in Schumacheria zygomorphic). Stamens more than 200 (rarely more than 900). Carpels usually numerous. Compitum usually absent. Exotestal cells in Dillenia large, tanniniferous, finally flattened.

Cladogram (simplified) of Dilleniaceae based on DNA sequence data (Horn 2009). Didesmandra and Neodillenia were not included in the analyses.

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PENTAPETALAE D. E. Soltis, P. S. Soltis et W. S. Judd

DILLENIACEAE Salisb.