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The families of flowering plants

L. Watson and M.J. Dallwitz

Phytolaccaceae R. Br.

Including Endochromaceae Dulac, Hilleriaceae Nakai, Lophiocarpaceae Doweld & Reveal, Microteaceae, Petivereae (Petiveriaceae) C.A. Agardh, Rivineae (Rivinaceae) J.G. Agardh, Seguiariaceae Nak.; excluding Achatocarpaceae, Agdestidaceae, Barbeuiaceae, Gisekiaceae Nak., 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. Leaf development not ‘graminaceous’. Domatia occurring in the family (found in Gallesia); manifested as hair tufts.

Leaf anatomy. The leaf lamina always with more than one layer of palisade, dorsiventral (usually), or bifacial (occasionaly isobilateral). Mucilaginous epidermis present, or absent. Stomata present; mainly confined to one surface, or on both surfaces; anomocytic, or paracytic. Hairs present, or absent (generally infrequent); eglandular; unicellular, or multicellular. Multicellular hairs uniseriate; simple. Complex hairs absent. Adaxial hypodermis present (occasionally), or absent. The mesophyll containing crystals, or without crystals. The crystals when present, raphides, or solitary-prismatic (styloids). Minor leaf veins without phloem transfer cells (Phytolacca, Rivina).

Axial (stem, wood) anatomy. Young stems with solid internodes to with hollow internodes. Pith with diaphragms, or without diaphragms. Cork cambium present; initially superficial. Nodes unilacunar. Primary vascular tissues comprising a ring of bundles, or comprising two or more rings of bundles; collateral. Internal phloem absent. Cortical bundles absent. Secondary thickening developing from a conventional cambial ring, or anomalous (commonly). The anomalous secondary thickening when present, via concentric cambia (resulting in successive zones of of vascular bundles embedded in secondary ground tissue). Primary medullary rays wide (generally lignified).

The vessels usually small. The vessel end-walls simple. The vessels usually without spiral thickening. The axial xylem with fibre tracheids, or without fibre tracheids; with libriform fibres, or without libriform fibres; including septate fibres (rarely), or without septate fibres. The fibres without spiral thickening. The parenchyma typically paratracheal. ‘Included’ phloem present (commonly, of the concentric type), or absent.

Reproductive type, pollination. Plants hermaphrodite (usually), or dioecious.

Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in ‘inflorescences’; when aggregated, in cymes, or in racemes, or in spikes, or in panicles. The ultimate inflorescence units 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; 3-celled (in several genera).

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. 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, phytochemistry. C3 (or CAM?). C3 physiology recorded directly in Phytolacca. Not cyanogenic. Alkaloids present, or absent. Iridoids not detected. Betalains present. Saponins/sapogenins present, or absent. Proanthocyanidins absent. Flavonols present, or absent; kaempferol. Ellagic acid absent (3 species, 3 genera). Aluminium accumulation not found. Plants accumulating free oxalates. Sieve-tube plastids P-type; type III (a).

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 III core angiosperms; Superorder Caryophyllanae. APG IV Order Caryophyllales.

Species about 100. Genera 15; Anisomeria, Ercilla, Gallesia, Hilleria, Ledenbergia, Lophiocarpus, Microtea, Monococcus, Petiveria, Phytolacca, Rivina, Schindleria, Seguieria, Trichostigma.

Economic uses, etc. A few cultivated as ornamentals.

Illustrations. • Monococcus echinophorus: Hook. Ic. Pl. 11 (1867–71). • Lophiocarpus polystachyus, as L. burchellii: Hook Ic. Pl. 15 (1884). • Petiveria alliacea: Hutchinson. • Phytolacca icosandra: Bot. Mag. 83 (1857). • Microtea debilis: Ann. Miss. Bot. Gard. 48 (1961). • floral diagrams of Mictotea maypurensis, Petiveria alliacea, Rivina humilis, Phytolacca spp.: Blütendiagramme 2 (1878). • Le Maout and Decaisne: Phytolacca, Seguieria. • Seguiera floribunda: Lindley. • Helleria secunda, as Rivina inaequalis: Hook. Ic. Pl. 2 (1837).

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: 5th March 2018.’.