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The families of flowering plants

L. Watson and M.J. Dallwitz

Flacourtiaceae DC.

~ Berberidopsidaceae

Including Bembiciaceae, Blackwelliaceae Sch. Bip., Caseariaceae F.N. Williams, Erythrospermaceae Van Tiegh., Homalineae (Homaliaceae) R.Br., Kiggelariaceae Link, Pangieae (Pangiaceae) Bl. ex Endl., Patrisiaceae Mart., Prockiaceae Bertuch, Samydaceae Ventenat, Soyauxiaceae Barkley; excluding Dioncophyllaceae, Gerrardinaceae, Neumanniaceae, Plagiopteraceae.

Habit and leaf form. Trees and shrubs; non-laticiferous, without coloured juice; leptocaul. Mesophytic. Leaves persistent; alternate; distichous (often), or spiral; ‘herbaceous’, or leathery; petiolate; non-sheathing; gland-dotted, or not gland-dotted; without marked odour; simple. Lamina entire; pinnately veined; cross-venulate. Leaves stipulate. Stipules caducous, or persistent. Leaf development not ‘graminaceous’. Domatia occurring in the family (known from 5 genera); manifested as pits, or hair tufts.

Leaf anatomy. The leaf lamina dorsiventral (usually), or centric. Hydathodes present, or absent. Epidermis with crystal idioblasts (very commonly), or without crystal idioblasts. Mucilaginous epidermis present, or absent. Stomata mainly confined to one surface (usually, abaxial), or on both surfaces; anomocytic, or paracytic. Hairs present (variously simple unicellular, tufted, 2-armed, stellate, peltate, uniseriate-eglandular or -glandular). Adaxial hypodermis present, or absent. Cystoliths present (Homalium), or absent. The mesophyll sometimes with secretory cavities containing resin; with sclerenchymatous idioblasts, or without sclerenchymatous idioblasts. Minor leaf veins without phloem transfer cells (Azara).

Axial (stem, wood) anatomy. Pith homogeneous, or heterogeneous. Cork cambium present; initially superficial. Nodes tri-lacunar. Primary vascular tissues in a cylinder, without separate bundles; collateral. Internal phloem absent. Cortical bundles absent. Medullary bundles absent. Secondary thickening developing from a conventional cambial ring. Primary medullary rays narrow (not widening in the phloem).

The wood diffuse porous. The vessels small, or medium, or large; solitary, or radially paired, or in radial multiples, or clustered. The vessel end-walls usually oblique; simple (usually), or scalariform and simple, or scalariform (rarely). The vessels without vestured pits; commonly with spiral thickening. The axial xylem with fibre tracheids, or without fibre tracheids; with libriform fibres, or without libriform fibres; including septate fibres (usually), or without septate fibres. The fibres without spiral thickening. The parenchyma when present, paratracheal. The secondary phloem not stratified. ‘Included’ phloem absent. The wood not storied. Tyloses absent.

Reproductive type, pollination. Plants hermaphrodite (usually), or monoecious (rarely), or dioecious (rarely). Pollination entomophilous.

Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in ‘inflorescences’; when aggregated, in cymes, or in fascicles, or in racemes, or in spikes, or in heads. The fruiting inflorescences conelike, or not conelike. The ultimate inflorescence units cymose (usually). Inflorescences terminal, or axillary, or epiphyllous (Bembicia, Phyllobotryon); pseudanthial (conelike, in Bembicia), or not pseudanthial. Flowers bracteate; bracteolate; small (usually), or medium-sized to very large (less often); regular; cyclic, or partially acyclic. When acyclic, the perianth acyclic, or the androecium acyclic, or the perianth acyclic and the androecium acyclic. Free hypanthium present (narrow), or absent. Hypogynous disk present; extrastaminal, or intrastaminal; of separate members, or annular (there being a disk, glands or scales between C and A or within A, and sometimes a ‘corona’ between C and A).

Perianth with distinct calyx and corolla, or sequentially intergrading from sepals to petals, or sepaline (corolla occasionally absent); 3–8, or 6–16(–30); when whorled 2 whorled (usually), or 3 whorled, or 1 whorled (rarely); when non-spiralled isomerous. Calyx 3–8(–15); when whorled 1 whorled; polysepalous, or gamosepalous; regular; persistent; accrescent, or non-accrescent; imbricate. Corolla 3–8(–15) (alternating with K, or spiral and intergrading with it); 1 whorled, or 2 whorled; appendiculate (sometimes with a ‘corona’ of scales inside, cf. Passifloraceae), or not appendiculate; imbricate; regular.

Androecium (4–)15–100 (i.e. usually ‘many’). Androecial members when many (i.e. usually), maturing centrifugally; free of the perianth; free of one another, or coherent; when united, tending to be 3–8 adelphous (in antepetalous groups); 1 whorled, or 2 whorled (or spiralled). Androecium exclusively of fertile stamens, or including staminodes. Stamens (4–)15–100 (usually ‘many’); isomerous with the perianth to diplostemonous to polystemonous. Anthers basifixed; non-versatile; dehiscing via longitudinal slits (usually), or dehiscing via pores (in Kiggelaria, these terminal); usually latrorse; tetrasporangiate; variously appendaged, or unappendaged. The anther appendages apical (often in the form of a prolonged connective). Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral. Anther wall initially with one middle layer, or initially with more than one middle layer. Pollen grains aperturate; mostly 3 aperturate; colporate (usually); 2-celled (only Casearia recorded).

Gynoecium 2–10 carpelled (with Aphloia excluded). The pistil 1 celled, or 2–10 celled. Gynoecium syncarpous (with Aphloia excluded); synovarious to synstylovarious (styles more or less united); superior (usually), or partly inferior (Bembicia). Ovary 1 locular (nearly always, but with the placentas more or less intruded), or 2–10 locular (effectively so, and genuinely so in e.g. Prockia). The ‘odd’ carpel when G3 posterior, or anterior. Styles partially joined; attenuate from the ovary. Stigmas (1–)2–10; dry type; non-papillate; Group II type. Placentation usually (i.e. when discernably unilocular), parietal; when plurilocular axile. Ovules in the single cavity when unilocular, 20–100 (i.e. ‘many’); 15–50 per locule (‘many’); pendulous; arillate, or non-arillate; orthotropous, or anatropous, or hemianatropous; bitegmic; crassinucellate. Outer integument contributing to the micropyle. Embryo-sac development Polygonum-type. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids hooked (sometimes with filiform apparatus). Hypostase present (Casearia), or absent. Endosperm formation nuclear.

Fruit fleshy, or non-fleshy; dehiscent, or indehiscent; a capsule, or a berry (usually), or a drupe. The drupes with separable pyrenes (Flacourtia). Seeds endospermic. Endosperm oily. Embryo well differentiated. Cotyledons 2; flat. Embryo achlorophyllous (1/2); straight (usually).

Seedling. Germination phanerocotylar, or cryptocotylar.

Physiology, phytochemistry. C3 (?), or CAM. CAM recorded directly in Flacourtia (non-succulent, and dubious). Sugars transported as sucrose, or as oligosaccharides + sucrose (but sucrose predominating, in all five genera sampled). Cyanogenic, or not cyanogenic. Cynogenic constituents of the gynocardin group. Alkaloids present, or absent. Iridoids not detected. Saponins/sapogenins present, or absent. Proanthocyanidins present (Azara), or absent; when present, cyanidin. Flavonols present, or absent; kaempferol, or quercetin, or kaempferol and quercetin. Ellagic acid absent (8 species, 7 genera). Aluminium accumulation demonstrated.

Geography, cytology. Sub-tropical to tropical. Pantropical. X = 10–12.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Violiflorae; Violales. Cronquist’s Subclass Dilleniidae; Violales. APG III core angiosperms; core eudicot; Superorder Rosanae; fabid. APG IV Order Malpighiales.

Species 1000. Genera about 90; Lately referred to Salicaceae sensu lato: Abatia (or Passifloraceae), Ahernia, Aphaerema (or Passifloraceae), Azara, Baileyoxylon, Banara (or Tiliaceae), Bartholomaea, Bembicia, Bennettiodendron, Byrsanthus, Calantica, Carpotriche, Carrierea, Casearia, Chlorocarpa, Dasylepis, Dissomera, Dovyalis, Eleutherandra, Euceraea, Flacourtia, Grandidiera, Hasseltia (or Tiliaceae), Hasseltiopsis (or Tiliaceae), Hecatostemon, Hemiscolopia, Homalium, Idesia, Itoa, Laetia, Lasiochlamys, Ludia, Lunania, Macrohasseltia (or Tiliaceae), Mayna, Mocquerysia, Neopringlea, Neoptychocarpus, Neosprucea (~ Tiliaceae), Olmediella, Oncoba, Ophiobotrys, Osmelia,Peterodendron, Phyllobotryon, Phylloclinium, Pineda (~ Tiliaceae), Pleuranthodendron, Poggea, Poliothyrsis, Priamosia, Prockia (or Tiliaceae), Prockiopsis, Pseudoscolopia, Pseudosmelia, Rawsonia, Ryania, Samyda, Scaphocalyx, Scolopia, Soyauxia, Tetrathylacium, Tisonia, Trichostephanus, Trimeria, Xylosma, Zuelania; lately referred to Achariaceae sensu lato: Buchnerodendron, Caloncoba, Camptostylus, Chiangiodendron, Erythrospermum, Gynocardia, Hydnocarpus, Kiggelaria, Lindackeria, Pangium, Ryparosa, Scottellia, Trichadenia, Xylotheca.

General remarks. This family, though universally accepted until quite recently, was evidently heterogeneous, and some of the constituent genera were referred to other families in earlier editions of this package. All the remaining genera (see the above lists) are now being referred to hugely enlarged versions of the formerly small families Salicaceae and Achariaceae; but it is of course much easier - but deplorable - to publish major realignments of genera than to organize functional, properly comparative descriptions of the kind required by serious scientific researchers ....

Economic uses, etc. Edible fruit (‘Ceylon gooseberry’) from Dovyalis.

Illustrations. • Azara dentata: Bot. Reg. 1788, 1836. • Azara petiolaris, as A. gilliesii: Bot. Mag. 86 (1860). • Bembicia axillaris: Hook. Ic. Pl. 15 (1883). • Byrsanthus brownii: Lindley. • Buchnerodendron lasiocalyx, as Oncoba: Hook Ic. Pl. 15 (1885). • Casearia grandiflora: Lindley. • Le Maout and Decaisne: Casearia, Samyda. • Dasylepis racemosa: Hook. Ic. Pl. 11 (1867–71). • Dovyalis hebecarpa: Trimen, Ill. Fl Ceylon (1893). • Erythrospermum hypoleucum: Hook. Ic. Pl. 19 (1889). • Erythrospermum acuminatissimum (as E. polyandrum): Hook. Ic. Pl. 14 (1880–82). • Euceraea nitida: Martius, Nova Gen. et Spec. Pl. Brasiliensium 3 (1829). • Flacourtia ramontchi: Thonner. • Homalium bionense, H. maringitra, H. stelliferum, H. vatkeanum: Humbert, Fl. de Madagascar et Comores 140 (1946). • Hydnocarpus inebrians: R. Wight (1840). • Idesia polycarpa: Bot. Mag. 111 (1885). • Le Maout and Decaisne: Pangium. • Le Maout and Decaisne: Pangium edule: Lindley. • Phyllobotryon spathulatum: Hook. Ic. Pl. 14 (1880–82). • Scottellia leonensis: Hook. Ic. Pl. 23 (1894). • Soyauxia gabonensis: Hook. Ic. Pl. 14 (1880–82). • Le Maout and Decaisne: Xylosma.

We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, genera included in each family, and classifications (Dahlgren; Dahlgren, Clifford, and Yeo; Cronquist; APG). See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 15th April 2018.’.