Phytolaccaceae R.Br.

Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://creativecommons.org/licenses/by/3.0

Morphology General Habit

Trees, woody climbers, shrubs or herbs (mostly perennial, sometimes annual), glabrous or with unbranched trichomes, these sometimes gland -tipped.  Leaves alternate, estipulate (but prophylls of axillary buds transformed into spines and resembling stipules in Seguieria Loefl.), usually petiolate, lamina simple, entire, pinnately veined.  Inflorescences mostly of spikes or racemes, rarely of panicles

Morphology Reproductive morphology Flowers

Flowers hermaphroditic or unisexual with rudiments of the opposite sex, actinomorphic or sometimes weakly zygomorphic (Anisomeria D. Don and Hilleria Vell); perianth in a single whorl, tepals 4 or 5, often persistent, imbricate, free or rarely connate at the base (Hilleria); stamens 3 to many, alternate or irregularly arranged with respect to the sepals, free, anthers bithecal, dorsifixed, introrose or rarely extrorse (Hilleria), with longitudinal dehiscence; ovary superior or rarely semi-inferior (Agdestis Mocino & Sessé ex DC), sessile or rarely on a gynophore (Nowickea J. Martínez & J. A. McDonald), 1-16-locular, locules connate or free, 1-carpellate ovaries with a single basal ovule, multi-carpellate ovaries with axillary placentation and a single ovule in each locule, campylotropous, styles usually free, rarely united or absent, stigmas free, capitate or penicellate

Morphology Reproductive morphology Fruits

Fruit a berry, samara, drupe or urticle; seeds 1 per locule, perisperm copious to lacking, embryo curved X=9.

Distribution

Agdestis Mocino & Sessé ex DC.: monotypic, occurring in tropical and subtropical habitats from the southeastern United States (Florida and Texas) to Central America; naturalized at least in the Antilles and Brazil. Anisomeria D. Don: three species endemic to Chile. Gallesia Casar.: monotypic, South America in Bolivia, Brazil, Ecuador, and Peru. Hilleria Vell.: three species, northern South America from Colombia and Venezuela to Bolivia and Brazil. Ledenbergia Klotzsch ex Moq.: two species, Mexico to South America. Microtea Sw.: approximately 10 species, Central America and the Antilles to South America. Nowickea J. Martínez & J. A. McDonald: two species endemic to central Mexico. Petiveria L.: monotypic, widespread in tropical and subtropical America from Florida and the Antilles to Argentina. Phytolacca L.: approximately 20 species, subcosmopolitan, nearly 16 species in the Neotropics. Rivinia L.: monotypic, widespread in tropical and subtropical America from the southern United States and the Antilles to Argentina and Chile; naturalized in the Old World. Schindleria H. Walter: two species in Peru and Bolivia. Seguieria Loefl.: 15-20 species distributed from Trinidad to Argentina with the majority of the species endemic to Brazil. Trichostigma A. Rich.: three species distributed from Mexico and the Antilles to Argentina. All of the genera present in the neotropics are native. Four are strictly endemic: Gallesia, Ledenbergia, Nowickea, and Schindleria.  Seven others extend out of the tropics proper into subtropical America: Agdestis, Anisomeria, Hilleria, Microtea, Petiveria, Seguieria, and Trichostigma. Only Phytolacca is also native to the Old World, but Rivinia is naturalized there.

Diagnostic

Key differences from similar families: Phytolaccaceae can be distinguished from most other Caryophyllalean families by the combination of indeterminate inflorescences (paniculate only in Gallesia and Seguieria) and either unicarpellate ovaries with a single basalovule or multicarpellate ovaries with axillary placentation and a single ovule in each locule. It is distinguished from Stegnospermataceae by indehiscent fruits (vs. capsules), the absence of petals (vs. petals present), and the lack of an aril on the seeds (vs. arillate seeds). It can be separated from Achatocarpaceae by having either bisexual flowers or unisexual flowers with rudiments of the opposite sex (vs. unisexual flowers lacking rudiments of the opposite sex) and uni- to multiovulate ovaries with the same number of ovules as carpels (vs. a uniovulate ovary formed by 2 carpels). Leaves petiolate. Flowers in spikes or racemes, actinomorphic, bisexual, with introrse anthers. Ovarysuperior, sessile. Distinguishing characters (always present): Leaves alternate, estipulate, with a simple, entire, pinnately veined lamina. Flowers in many-flowered inflorescences. Perianth in a single whorl of 4 or 5 imbricate tepals. Fruitindehiscent with 1 seed per locule. Key to genera of Neotropical Phytolacaceae 1. Leaves strongly cordate at the base; ovary partially inferior — Agdestis1. Leaves not cordate at the base; ovary superior... 2 2. Ovary of 5-16 carpels and seeds ... 32. Ovary of a single carpel and seed... 5 3. Sepals unequal and somewhat fleshy; carpels distinctly free — Anisomeria3. Sepals subequal and not fleshy; carpels (at least in the neotropical species) united ... 4 4. Ovary on an elongated gynophore — Nowickea 4. Ovary sessile, gynophore absent — Phytolacca 5. Ovary with 2-4 stigmas — Microtea5. Ovary with a single stigma... 6 6. Flowers slightly zygomorphic, upper tepalfree, the lower three tepals connate at the base into a three-lobed lip — Hilleria6. Flowers actinomorphic, all tepals free... 7 7. Ovary with 4-6 recurved, awn -like protuberances — Petiveria7. Ovary without protuberances ... 8 8. Inflorescence paniculate; fruit a samara ... 98. Inflorescence spiciform or racemose; fruit a drupe or utricle ... 10 9. Plants usually spiny; tepals 5, herbaceous and reflexed in fruit — Seguieria9. Plants unarmed; tepals 4, woody and erect in fruit — Gallesia 10. Fruit a drupe... 1110. Fruit a utricle ... 12 11. Stamens 4; stigma capitate — Rivinia11. Stamens 8-25; stigma penicillate — Trichostigma 12. Racemes pendulous; bract placed halfway or above between the flower and the base of the pedicel; tepals reticulate veined — Ledenbergia12. Racemes erect; bract placed at the base of the pedicel; tepals parallel veined — Schindleria

Note

Number of genera: The family contains approximately 16 genera and 75-85 species. Nearly 60 species in 13 genera are present in the Neotropics: Agdestis, Anisomeria, Gallesia, Hilleria, Ledenbergia, Microtea, Nowickea, Petiveria, Phytolacca, Rivinia, Schindleria, Seguieria, and Trichostigma. The greatest diversity of genera and species occurs in the Neotropics, particularly South America. The majority of taxa are herbaceous and most of the woody members have anomalous secondary thickening by means of successive cambia. All of the American genera are present in the Neotropics except Ercilla Adr. Juss. (1 or 2 spp.) which occurs in mesic temperate forests of Chile. Two additional genera are restricted to the Old World: Lophiocarpus Turcz. and Monococcus F. Muell. The family is of little economic importance.  Phytolaccadioica L., ombú tree, is occasionally cultivated in warmer regions of the world, mostly as a fast-growing shade tree having a peculiar trunk and soft, spongy wood.  Petiveria and some species of Gallesia and Phytolacca are used in traditional medicine.  The high saponin content of some species makes them useful for washing clothes, and in eastern Africa Phytolaccadodecandra L'Hér., endod or soap berry plant, is employed locally for this purpose.  This species also has molluscicide properties and is used to control schistosomiasis. The largest and most taxonomically difficult genus is Phytolacca in which species boundaries are often blurred due to widespread hybridization. For a relatively small family, fruit variation is notable with berries, drupes, utricles and samaras present. Notes on delimitation: Although Phytolaccaceae clearly belongs to the Caryophyllales, there is much debate about its circumscription and exact position.  Some genera historically considered within the family have recently been segregated as separate families.  For example, the North American genus Stegnosperma Benth. is now considered in a monogeneric family Stegnospermataceae, as too are the Old World genera Barbeuia Thouars. (Barbeuiaceae) and Gisekia L. (Gisekiaceae). The New World genera Achatocarpus Triana. and Phaulothamnus A. Gray were recognized by some specialists as members of Phytolaccaceae, but these are now treated in Achatocarpaceae.  Even with these genera removed, controversy exists, and the core of the family is sometimes further divided into two families: Phytolaccaceae (ovary of 3-16 carpels, corresponding to subfamiles Agdestioideae and Phytolaccoideae) and Petiveriaceae (ovary of one carpel, corresponding to subfamilies Rivinioideae and Microteoideae). A definitive conclusion as to their status awaits further comprehensive studies, and the recommendations for a broad Phytolaccaceae are followed here.

Phytolaccaceae, R. M. Polhill. Flora of Tropical East Africa. 1971

Morphology General Habit

Woody or herbaceous, erect or scrambling

Morphology Leaves

Leaves alternate, simple, entire, pinnately nerved, with crystals generally easily visible in the younger leaves

Morphology Leaves Stipules

Stipules generally lacking

Morphology Reproductive morphology Inflorescences

Inflorescences racemose, spicate or paniculate, terminal, lateral or axillary; bracts and paired bracteoles small

Morphology Reproductive morphology Flowers

Flowers small, hermaphrodite or ?and ? (then rudiments at least of aborted organs still present), usually regular

Morphology Reproductive morphology Flowers Calyx

Sepals 4–5 (rarely more), free or partially united, imbricate, usually persistent, sometimes brightly coloured

Morphology Reproductive morphology Flowers Corolla

Petals generally absent (present only in Stegnosperma)

Morphology Reproductive morphology Flowers Androecium

Stamens (3–)4–many in 1–several whorls, when definite alternate with the sepals, usually inserted on a ± fleshy disk; filaments free or united at the base; anthers 2-thecous, dehiscing lengthwise

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior or rarely semi-inferior, of 1–many free or united 1-ovulate carpels; styles as many as the carpels, free or connate at the base, short, sometimes lacking; stigmas linear or capitate; ovule basal, shortly stalked, campylotropous

Morphology Reproductive morphology Fruits

Fruit of 1–numerous free or united carpels, fleshy or dry, sometimes winged

Morphology Reproductive morphology Seeds

Seed subglobose, discoidal or reniform, with a membranous or brittle testa, occasionally arillate; embryo annular, surrounding the endosperm

Phytolaccaceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:1. 1954

Morphology General Habit

Herbs, shrubs or trees

Morphology Leaves

Leaves alternate, entire; stipules absent or minute

Morphology Reproductive morphology Flowers

Flowers hermaphrodite or unisexual, actinomorphic or nearly so, hypogynous, in simple or compound terminal or axillary racemes

Morphology Reproductive morphology Flowers Perianth

Perianth simple, herbaceous or coriaceous

Morphology Reproductive morphology Flowers Calyx

Sepals free or some connate, imbricate, equal or unequal

Morphology Reproductive morphology Flowers Corolla

Petals absent

Morphology Reproductive morphology Flowers Androecium

Stamens 3 to many, often inserted on a hypogynous disk; filaments free or slightly connate at the base; anthers 2-celled, cells parallel and dehiscing longitudinally

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, composed of 1 or several carpels either free or variously connate; style almost none or short; ovule solitary in each carpel, basal

Morphology Reproductive morphology Fruits

Fruit of 1 or many carpels, fleshy or dry

Morphology Reproductive morphology Seeds

Seed erect, with the large embryo in the periphery and enclosing the endosperm

George R. Proctor (2012). Flora of the Cayman Isands (Second Edition). Royal Botanic Gardens, Kew

Morphology General Habit

Herbs, shrubs or trees, mostly glabrous; leaves alternate, simple, entire; stipules minute or lacking

Morphology Reproductive morphology Flowers

Flowers regular, perfect, in terminal and axillary racemes or spikes; perianth of 4 or 5 segments, persistent in fruit

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens 4, 5, or more, variously inserted

Morphology Reproductive morphology Flowers Gynoecium Ovary

Ovary superior, of 1–several 1-ovuled carpels, the ovules basal; style short or lacking

Morphology Reproductive morphology Fruits

Fruit an achene or berry of 1–several carpels; seeds with mealy or fleshy endosperm.

Distribution

A widespread, chiefly tropical family of up to 22 genera and about 120 species.