1. Phytolaccaceae R.Br.

    1. This family is accepted.

[FZ]

Phytolaccaceae, B. L. Stannard. Flora Zambesiaca 9:1. 1988

Habit
Trees, shrubs or herbs, sometimes scrambling
Leaves
Leaves simple, alternate, entire, petiolate to sessile, crystals often visible especially on younger leaves
Stipules
Stipules absent (in the Flora Zambesiaca area), minute or thorny
Inflorescences
Inflorescences terminal or axillary, spicate, racemose or paniculate
Flowers
Flowers bracteate and bracteolate, hermaphrodite or unisexual (then usually with rudimentary aborted organs), usually actinomorphic
Calyx
Sepals 4–5, free or connate towards base, imbricate
Corolla
Petals absent
Androecium
Stamens (3) 4-many, 1–2-seriate, often inserted on more or less fleshy, annular disk, irregularly arranged or alternate to sepals; filaments slender, free or connate at base; anthers dorsi-or basifixed, dehiscing longitudinally
Gynoecium
Ovary usually superior, composed of 1-many free or connate carpels; ovule basal, solitary in each carpel; styles absent or same number as carpels, free or united at base; stigmas linear or capitate
Fruits
Fruit of 1-many, free or connate carpels, fleshy or dry, sometimes winged; seed subglobose, discoidal or reniform, often compressed; testa membranous or brittle; endosperm present
[NTK]

Steinmann, V.W. (2010). Neotropical Phytolaccaceae.

Morphology
Description

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

Distribution
Distribution in the Neotropics
  • 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.
Diagnostic
Notable genera and distinguishing features
  • See above
Useful tips for generic identification

Key to genera of Neotropical Phytolacaceae

1. Leaves strongly cordate at the base; ovary partially inferior — Agdestis
1. Leaves not cordate at the base; ovary superior... 2

2. Ovary of 5-16 carpels and seeds ... 3
2. Ovary of a single carpel and seed... 5

3. Sepals unequal and somewhat fleshy; carpels distinctly free — Anisomeria
3. 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 — Microtea
5. 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 — Hilleria
6. Flowers actinomorphic, all tepals free... 7

7. Ovary with 4-6 recurved, awn -like protuberances — Petiveria
7. Ovary without protuberances ... 8

8. Inflorescence paniculate; fruit a samara ... 9
8. Inflorescence spiciform or racemose; fruit a drupe or utricle ... 10

9. Plants usually spiny; tepals 5, herbaceous and reflexed in fruit — Seguieria
9. Plants unarmed; tepals 4, woody and erect in fruit — Gallesia

10. Fruit a drupe... 11
10. Fruit a utricle ... 12

11. Stamens 4; stigma capitate — Rivinia
11. 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 — Ledenbergia
12. Racemes erect; bract placed at the base of the pedicel; tepals parallel veined — Schindleria

Other important characters
  • 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 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).
General Description
General notes
  • 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.
Status
  • 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.
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.
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.
Literature
Important literature

Burger, W. 1983. Phytolaccaceae, Family 66. In: W. Burger (ed.). Flora Costaricensis. Fieldiana Bot. 13: 199-212.

Elliasson, U.H. 1993. Phytolaccaceae, pp 1-43. In: G. Harling and L. Andersson (eds). Flora of Ecuador vol. 35A. University of Gothenburg, Gothenburg.

Lozada, L. 2000. Phytolaccaceae. In: N. Diego-Pérez and R.M. Fonseca (eds.). Flora de Guerrero 10: 1-20.

Martínez-García, J. 1984. Phytolaccaceae. In: A. Goméz-Pompa (ed.). Flora de Veracruz 36: 1-41.

Martínez G., J. and McDonald, J.A. 1989. Nowickea (Phytolaccaceae), a new genus with two new species from Mexico. Brittonia 41: 399-403.

Nowicke, J.W. 1968. Palynotaxonomic study of the Phytolaccaceae. Ann. Missouri Bot. Gard. 55: 294-364.

Rogers G.K. 1985. The genera of Phytolaccaceae in the Southeastern United States-J. Arnold Arbor. 66: 1-37.

Rohwer, J.G. 1982. A taxonomic revision of the genera Seguieria Loefl. and Gallesia Casar. (Phytolaccaceae). Mitt. Bot. Staatssamml. München 18: 231-288.

Rohwer, J.G. 1993. Phytolaccaceae. Pp. 506-515. In: K. Kubitzki, J.G. Rohwer and V. Bittrich (eds.). The families and genera of vascular plants. Volume 2. Flowering plants. Springer-Verlag, Berlin.

Rzedowski, J. and Calderón de Rzedowski, G. 2000. In: J. Rzedowski and G. Calderón de Rzedowski (eds.). Flora del Bajío y de Regiones Adyacentes 91: 1-32.

Walter, H. 1906. Die Diagramme der Phytolaccacean. Bot. Jahrb. 37 (Beibl. 85): 1-57.

Walter, H. 1909. Phytolaccaceae. In: A. Engler (ed.). Pflanzenreich IV 83 (Heft 39): 1-154.

Walter, H. 1909. Phytolaccaceae. In: A. Engler (ed.). Pflanzenreich IV 83 (Heft 39): 1-154.

[FTEA]

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

Habit
Woody or herbaceous, erect or scrambling
Leaves
Leaves alternate, simple, entire, pinnately nerved, with crystals generally easily visible in the younger leaves
Stipules
Stipules generally lacking
Inflorescences
Inflorescences racemose, spicate or paniculate, terminal, lateral or axillary; bracts and paired bracteoles small
Flowers
Flowers small, hermaphrodite or ?and ? (then rudiments at least of aborted organs still present), usually regular
Calyx
Sepals 4–5 (rarely more), free or partially united, imbricate, usually persistent, sometimes brightly coloured
Corolla
Petals generally absent (present only in Stegnosperma)
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
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
Fruits
Fruit of 1–numerous free or united carpels, fleshy or dry, sometimes winged
Seeds
Seed subglobose, discoidal or reniform, with a membranous or brittle testa, occasionally arillate; embryo annular, surrounding the endosperm
[FWTA]

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

Habit
Herbs, shrubs or trees
Leaves
Leaves alternate, entire; stipules absent or minute
Flowers
Flowers hermaphrodite or unisexual, actinomorphic or nearly so, hypogynous, in simple or compound terminal or axillary racemes
Perianth
Perianth simple, herbaceous or coriaceous
Calyx
Sepals free or some connate, imbricate, equal or unequal
Corolla
Petals absent
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
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
Fruits
Fruit of 1 or many carpels, fleshy or dry
Seeds
Seed erect, with the large embryo in the periphery and enclosing the endosperm

Images

Phytolaccaceae R.Br. appears in other Kew resources:

First published in Narr. Exped. Zaire 454. 1818 [5 Mar 1818] (1818)

Accepted by

  • APG IV (2016) http://dx.doi.org/10.1111/boj.12385

Sources

Flora Zambesiaca
Flora Zambesiaca
http://creativecommons.org/licenses/by-nc-sa/3.0

Flora of Tropical East Africa
Flora of Tropical East Africa
http://creativecommons.org/licenses/by-nc-sa/3.0

Flora of West Tropical Africa
Flora of West Tropical Africa
http://creativecommons.org/licenses/by-nc-sa/3.0

Kew Names and Taxonomic Backbone
The International Plant Names Index and World Checklist of Selected Plant Families 2019. Published on the Internet at http://www.ipni.org and http://apps.kew.org/wcsp/
© Copyright 2017 International Plant Names Index and World Checklist of Selected Plant Families. http://creativecommons.org/licenses/by/3.0

Neotropikey
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