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 BarkleyExcluding Dioncophyllaceae, Gerrardinaceae, Neumanniaceae, Plagiopteraceae
Habit and leaf form. Trees and shrubs, or lianas (Berberidopsis); non-laticiferous and without coloured juice. Self supporting. Leptocaul. Mesophytic. Leaves evergreen; 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. Leaves without a persistent basal meristem. Domatia occurring in the family (from 5 genera); manifested as pits, or hair tufts.
Leaf anatomy. Epidermis very commonly with crystal idioblasts. Mucilaginous epidermis present, or absent. Stomata anomocytic, or paracytic.
Adaxial hypodermis present, or absent. Lamina dorsiventral (usually), or centric. Cystoliths present (Homalium), or absent. The mesophyll with sclerencymatous idioblasts, or without sclerenchymatous idioblasts. Minor leaf veins without phloem transfer cells (Azara, Berberidopsis).
Stem anatomy. Cork cambium present; initially superficial. Nodes tri-lacunar. Primary vascular tissue in a cylinder, without separate bundles. Cortical bundles absent. Medullary bundles absent. Internal phloem absent. Secondary thickening developing from a conventional cambial ring. The secondary phloem not stratified. ‘Included’ phloem absent. Xylem with fibre tracheids, or without fibre tracheids; with libriform fibres, or without libriform fibres. Vessel end-walls simple, or scalariform and simple, or scalariform (rarely). Vessels without vestured pits. Wood parenchyma when present, paratracheal.
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’; in cymes, in fascicles, in racemes, in spikes, and in heads. The fruiting inflorescences conelike, or not conelike. The ultimate inflorescence unit cymose (usually). Inflorescences terminal, or axillary, or epiphyllous (Bembicia); 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, biochemistry. Cyanogenic, or not cyanogenic. Cynogenic constituents of the gynocardin group. Alkaloids present, or absent. Iridoids not detected. Proanthocyanidins present (Azara, Berberidopsis), or absent; when present, cyanidin. Flavonols present, or absent; kaempferol, or quercetin, or kaempferol and quercetin. Ellagic acid absent (8 species, 7 genera). Saponins/sapogenins present, or absent. Aluminium accumulation demonstrated. Sugars transported as sucrose, or as oligosaccharides + sucrose (but sucrose predominating, in all five genera sampled). C3 (?), or CAM. CAM recorded directly in Flacourtia (non-succulent, and dubious).
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 3 core angiosperms; core eudicot; Superorder Rosanae; fabid; 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, Trichadenia, Xylotheca,.
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 to publish major realignments of genera than to organize functional, properly comparative descriptions of the kind required here ....
Economic uses, etc. Edible fruit (‘Ceylon gooseberry’) from Dovyalis.
• Technical details: Byrsanthus (Lindley).
• Technical details: Casearia (Lindley).
• Technical details: Casearia, Samyda.
• Technical details: Flacourtia (Thonner).
• Technical details: (Pangium).
• Technical details: Pangium (Lindley).
• Technical details: (Xylosma).
• Azara dentata: Bot. Reg. 1788, 1836.