Phytolaccaceae R. Br.
Including Endochromaceae Dulac, Gisekiaceae Nak., Hilleriaceae Nakai, Petivereae (Petiveriaceae) C.A. Agardh, Rivineae (Rivinaceae) J.G. Agardh, Seguiariaceae Nak.
Excluding Achatocarpaceae, Agdestidaceae, Barbeuiaceae, Stegnospermataceae
Habit and leaf form. Herbs (usually), or trees, or shrubs, or lianas. Plants more or less succulent, or non-succulent. Self supporting (usually), or climbing. Mesophytic, or xerophytic. Leaves alternate; spiral; ‘herbaceous’, or fleshy; petiolate to sessile; non-sheathing; gland-dotted, or not gland-dotted; simple. Lamina entire; pinnately veined; cross-venulate. Leaves stipulate (e.g. Seguieria), or exstipulate (usually). Stipules when present, free of one another; spiny. Lamina margins entire. Leaves without a persistent basal meristem. Domatia recorded (Gallesia); represented by hair tufts.
Leaf anatomy. Mucilaginous epidermis present, or absent.
Minor leaf veins without phloem transfer cells (Phytolacca, Rivina).
Stem anatomy. Cork cambium present; initially superficial. Nodes unilacunar. Internal phloem absent. Secondary thickening developing from a conventional cambial ring, or anomalous; when anomalous, via concentric cambia. ‘Included’ phloem present (often?), or absent. Xylem with fibre tracheids; with libriform fibres, or without libriform fibres; with vessels. Vessel end-walls simple. Sieve-tube plastids P-type; type III (a). Pith with diaphragms, or without diaphragms.
Reproductive type, pollination. Plants hermaphrodite (usually), or dioecious.
Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in ‘inflorescences’; in cymes, or in racemes, or in spikes, or in panicles. The terminal inflorescence unit cymose, or racemose. Inflorescences terminal, or axillary, or leaf-opposed; spikes to panicles, rarely cymes. Flowers bracteate; bracteolate (the bracts and bracteoles small); small; regular (variable in form, showing unusual diversity in androecium and gynoecium); cyclic. Floral receptacle developing a gynophore, or with neither androphore nor gynophore. Free hypanthium absent. Hypogynous disk present, or absent.
Perianth sepaline (corolla absent, the calyx green or somewhat coloured); 4, or 5(–10); 1 whorled. Calyx 4, or 5(–10) (the segments equal or unequal); 1 whorled; usually persistent; imbricate.
Androecium 4–5, or 5–100 (i.e. to ‘many’). Androecial members branched, or unbranched; when branched/many, maturing centrifugally; free of the perianth; free of one another, or coherent (then the filaments joined basally); when connate, 1 adelphous; 1 whorled, or 2 whorled. Androecium exclusively of fertile stamens. Stamens 4–5, or 5–50; isomerous with the perianth, or diplostemonous to polystemonous. Anthers dorsifixed; dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. Anther wall of the ‘monocot’ type. Tapetum glandular. Pollen grains aperturate; 3–12(–30) aperturate; colpate, or foraminate, or rugate; spinulose; 2-celled, or 3-celled (usually?).
Gynoecium (1–)4–12(–16) carpelled. Carpels isomerous with the perianth, or reduced in number relative to the perianth, or increased in number relative to the perianth. The pistil when other than apocarpous, 2–16 celled (?). Gynoecium monomerous (rarely), or apocarpous, or syncarpous; of one carpel to eu-apocarpous, or semicarpous to synovarious (i.e. the carpels free to more or less connate); superior. Carpel when apocarpous/semicarpous, stylate; 1 ovuled. Placentation when apocarpous, basal. Ovary when more or less syncarpous, 2–16 locular (?). Styles 2–16 (?); free, or partially joined. Stigmas 2–16. Placentation when more or less syncarpous, basal. Ovules 1 per locule; arillate, or non-arillate; campylotropous, or amphitropous; bitegmic; crassinucellate. Outer integument not contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; proliferating (Rivina, up to 6 cells), or not proliferating; ephemeral, or persistent. Synergids pear-shaped, or hooked (with filiform apparatus, in Gisekia). Endosperm formation nuclear. Embryogeny onagrad, or caryophyllad, or chenopodiad.
Fruit fleshy, or non-fleshy; an aggregate, or not an aggregate. The fruiting carpel when apocarpous, indehiscent; samaroid, or nucular, or drupaceous. Fruit when syncarpous, dehiscent, or indehiscent, or a schizocarp (when G2). Mericarps when schizocarpic, 2; comprising berrylets, or comprising nutlets, or comprising drupelets. Fruit when more or less syncarpous, a capsule, or a berry. Capsules loculicidal. Seeds non-endospermic. Perisperm present. Seeds with starch. Embryo well differentiated. Cotyledons flat, or folded. Embryo achlorophyllous (2/2); curved. Micropyle not zigzag.
Seedling. Germination phanerocotylar.
Physiology, biochemistry. Not cyanogenic. Alkaloids present, or absent. Iridoids not detected. Proanthocyanidins absent. Flavonols present, or absent; kaempferol. Ellagic acid absent (3 species, 3 genera). Betalains present. Saponins/sapogenins present, or absent. Aluminium accumulation not found. Plants accumulating free oxalates. C3. C3 physiology recorded directly in Phytolacca.
Geography, cytology. Temperate to tropical. Tropical and warm temperate America, Africa, Eurasia, Southeast Asia, Australia. X = 9.
Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Caryophylliflorae; Caryophyllales. Cronquist’s Subclass Caryophyllidae; Caryophyllales. APG (1998) Eudicot; core Eudicot; neither Rosid nor Asterid; Caryophyllales. Species about 100. Genera 15; Anisomeria, Ercilla, Gallesia, Gisekia, Hilleria, Ledenbergia, Lophiocarpus, Microtea, Monococcus, Petiveria, Phytolacca, Rivina, Schindleria, Seguieria, Trichostigma.
Economic uses, etc. A few cultivated as ornamentals.
Illustrations. • Technical details (Phytolacca, Seguiera). • Technical details (Seguiera).
Cite this publication as: ‘L. Watson and M. J. Dallwitz (1992 onwards). The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 14th December 2000. http://biodiversity.uno.edu/delta/’. Dallwitz (1980), Dallwitz, Paine and Zurcher (1993, 1995, 2000), and Watson and Dallwitz (1991) should also be cited (see References).
