1. Balanophoraceae Rich.

    1. This family is accepted.

[FWTA]

Balanophoraceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:2. 1958

Habit
Fleshy herbs parasitic on roots, annual or perennial, destitute of chlorophyll and stomata
Flowers
Flowers unisexual, very rarely hermaphrodite, densely crowded into unisexual or androgynous inflorescences; male flowers without or with a valvate 3–8-lobed perianth
Androecium
Stamens 1–2 in the achlamydeous flowers, in those with a perianth often equal in number to, and opposite the lobes; filaments free or connate; anthers 2–4-celled or with many cells, free or connate, opening by pores or slits
Gynoecium
Ovary 1–3-celled, adnate to the perianth when present; styles 1–2, terminal or rarely the stigma sessile and discoid; ovule solitary in each cell, mostly pendulous, nude or with a single integument
Fruits
Fruit small, nut-like, 1-celled, 1-seeded
Seeds
Seeds with abundant endosperm and very small embryo
[NTK]

Delprete, P.G. (2009). Neotropical Balanophoraceae.

Morphology
Description

Herbaceous , fleshy plants, entirely or partially subterraneous, "fungus-like", parasitic on roots of trees, shrubs or herbs, without chlorophyll, without stomata , with white, yellow, red or brown color; " tuber " (undefined root / stem structure) branched, unbranched, globose , or irregularly shaped, with the inflorescence originating directly from the " tuber ". Leaves reduced to scales, spirally arranged, margin entire , venation not evident. Inflorescence spicate or spadix -shaped, the lateral branches much reduced or absent; bracts scale-like, triangular, ovate , hexagonal or almost round, peltate . Flowers unisexual. Male flowers 3- merous , with 3- lobed perianth , or a synandrium of 3 free anthers; anthers 2(3)-locular, basifixed, opening by longitudinal slit. Female flowers with a 2- lobed or irregularly- lobed perianth , or absent; ovary inferior; ovary without well-defined locules , placenta , and ovules; styles 1 or 2, stigma minutely capitate . Fruit a minute, 1-seeded achene .

Distribution
Distribution in the Neotropics
  • Corynaea Hook.f. (1 sp.): Panama, Colombia, Venezuela, Ecuador, Peru, and Bolivia.
  • Helosis Rich. (1 sp.): Mexico, Central America, Greater Antilles, Colombia, Ecuador, Peru, Bolivia, Venezuela, Guianas, throughout southern Brazil.
  • Langsdorffia Mart. (1 sp.): Throughout the Neotropics, from Mexico to southern Brazil.
  • Lathrophytum Eichl. (1 sp.): Brazil (states of Goiás and Rio de Janeiro).
  • Lophophytum Schott & Endl. (4 spp.): Colombia, Peru, Bolivia, Brazil, Paraguay, and northern Argentina.
  • Ombrophytum Poepp. ex Endl. (4 spp.): Ecuador, Galapagos Islands, Peru, Bolivia, Brazil, Chile, and Argentina.
  • Scybalium Schott & Endl. (4 spp.): Greater Antilles, Colombia, Venezuela, Ecuador, and Brazil.
Diagnostic
Key differences from similar families
  • Family very distinctive from other parasitic taxa, by its set of unique characters.
  • The other family of subterranean holoparasitic root parasites with subterranean habit is the Hydnoraceae, which is a characterized by a large, fleshy, solitary flower, with 3-4 fleshy tepals, and a large fruit with numerous seeds.
Useful tips for generic identification

Key to genera of Neotropical Balanophoraceae

Key adapted from Hansen (1980):

1. Style 1; tubers with wax; flowers not embedded in filiform hairs, or anthers 3, merged into a synandrium ... Langsdorffia
1. Styles 2; tubers with starch; flowers with both sexes imbedded in filiform hairs, or anthers 2, not merged into a synandrium ... 2 

2. Flowers embedded in a layer of filiform hairs; anthers merged into a usually 3-merous  synandrium ... 3
2. Flowers not imbedded in hairs, on conspicuous or elongated branches, these subtended by deciduous scale-like bracts; stamens with filaments ... 5

3. Stem and young inflorescence covered by triangular scales ... Scybalium
3. Stem naked, or with a few inconspicuous bracts; young inflorescence covered by hexagonal, peltate bracts ... 4 

4. Inflorescence emerging from buds on elongated, rhizome-like structures, these appearing from a "tuber" connected with the host root... Helosis
4. Inflorescence emerging directly from a "tuber" connected with the host root... Corynaea

5. Inflorescence branches subtended by ovate -triangular bracts; apical part of female branch not peltately enlarged ... Lophophytum
5. Inflorescence branches subtended by hexagonal or irregularly-shaped bracts; apical part of female branch peltately enlarged, covering the flowers ... 6  

6. Male flowers solitary, each flower subtended by a peltatebract; anthers sessile... Lathrophytum
6. Male flowers in many-flowered branches, these subtended by a peltate or clavate bract; stamens with evident filaments ... Ombrophytum

Notable genera and distinguishing features

See above

Distinguishing characters (always present)
  • Parasitic plants, on roots trees, shrubs or herbs.
  • Entirely or partially subterraneous.
  • Without chlorophyll.
  • Without stomata.
  • White, yellow, red or brown color.
  • "Tuber" branched, unbranched, globose, or irregularly shaped.
  • Inflorescence originating directly from the "tuber ".
  • Flowers minute, unisexual,: male flowers 3-merous, female flowers 2-merous.
  • Ovary without well-defined locules, placenta, and ovules.
  • Fruit a minute, 1-seeded achene.
General Description
Number of genera

Worldwide 18 genera and 45 species. In the Neotropics, 7 genera and 16 species:

  • Corynaea (1 sp.)
  • Helosis (1 sp.)
  • Langsdorffia (1 sp)
  • Lathrophytum (1 sp.)
  • Lophophytum (4 spp.)
  • Ombrophytum (4 spp.)
  • Scybalium (4 spp.)
Status
  • A poorly known family in terms of distribution, pollination, and number of species, mostly due to their partially or entirely subterranean habit.
  • Some reports of plants visited by flies, bees and wasps, but no definitive studies about their pollination biology. Also, some reports about seed dispersal by ants.
  • Their general physiology is also little-known, as is a family without stomata.
Literature
Important literature

Borchsenius, F. & , J.M. Olesen. 1990. The Amazonian root holoparasite Lophophytum mirabile (Balanophoraceae) and its pollinators and herbivores. J. Trop. Ecol. 6: 501-505.

Delprete, P.G. 2004. A new species of Lophophytum and the first report of Lathrophytum (Balanophoraceae) from the state of Goiás, Central Brazil. Kew Bull. 59: 291-295.

Hansen, B. 1980. Balanophoraceae. Fl. Neotrop. Monogr. 23: 1-80.

Kuijt, J. & W.X. Dong. 1990. Surface features of the leaves of Balanophoraceae - A family without stomata? Pl. Syst. Evol. 170: 29-35.

[FTEA]

Balanophoraceae, B. Hansen. Flora of Tropical East Africa. 1993

Habit
Herbaceous, fleshy root-parasites, without chlorophyll and roots, yellowish white to yellow, orange to red or brown to black
Tubers
Tuber cylindrical or subspherical, branched or unbranched, irregularly corrugated or variously polygonately furrowed or ± warty, subterranean, developing by germination of a fruit on a suitable host root, producing one or more inflorescence-bearing stems
Stem
Stems with or without leaves, unbranched
Leaves
Leaves scaly, spirally arranged or distichous, decussate or verticillate, sometimes much reduced
Inflorescences
Inflorescence branched or appearing spadix-like, the branches much suppressed, rarely composed of minute cymes (Cynomorium L.); branches of first order bracteate, well-developed to totally suppressed, sometimes with the sterile apical part peltately widened; bracts scaly, ± transformed and reduced, triangular or peltate or reduced to a clavate structure, caducous or persistent, Flowers unisexual, rarely bisexual
Flowers
Male flowers without a perianth, or perianth of 3–7 segments Female flowers without a perianth or with a superior perianth inconspicuously 2-lobed or rarely 3–6-lobed
Male
Male flowers without a perianth, or perianth of 3–7 segments
Androecium
Stamens 1–7 or united into a synandrium, opposite the perianth-segments; anthers dehiscing longitudinally or irregularly
Female
Female flowers without a perianth or with a superior perianth inconspicuously 2-lobed or rarely 3–6-lobed
Gynoecium
Ovary 1-locular or without a cavity and any definite placenta; styles 1–2(–3), alternate with perianth-lobes; stigma inconspicuous or ± capitellate; generally one few-celled embryo developing in the central tissue of the ovary, surrounded by a few-celled endosperm and a layer of stone cells at maturity
Fruits
Fruit a small 1-seeded achene
[FZ]

Flora Zambesiaca. Vol. 9, Part 3. Polygonaceae-Myriaceae. Pope GV, Polhill RM, Martins ES. 2006.

Habit
Herbaceous parasites on roots of woody plants, without chlorophyll, usually brightly coloured, fleshy, arising from underground tubers attached to host plant, monoecious, dioecious or polygamous Herbaceous parasites on roots of woody plants, without chlorophyll, usually brightly coloured, fleshy, arising from underground tubers attached to host plant, monoecious, dioecious or polygamous
Tubers
Tubers cylindrical or subspherical, irregularly lobed or emitting branched cylindrical rhizomes Tubers cylindrical or subspherical, irregularly lobed or emitting branched cylindrical rhizomes
Inflorescences
Inflorescences simple or branched, with or without bracts Inflorescences simple or branched, with or without bracts
Flowers
Flowers borne in involucrate heads or in dense fleshy inflorescences Female flowers: perianth absent or much reduced, 3–several-lobed and adnate to ovary Male flowers: perianth segments (tepals) 3–9; stamens 2–12, free or united into a synandrium Flowers borne in involucrate heads or in dense fleshy inflorescences Female flowers: perianth absent or much reduced, 3–several-lobed and adnate to ovary Male flowers: perianth segments (tepals) 3–9; stamens 2–12, free or united into a synandrium
Ovary
Ovary inferior, 1–3-locular, ovules 1–3
Style
Style 1, long, filiform, or absent
Fruits
Fruit indehiscent, aggregated into syncarpia or separate. Fruit indehiscent, aggregated into syncarpia or separate
Note
I have largely followed B. Hansen's treatment in his ‘Balanophoraceae of Continental Africa’ (Bot. Jahrb. Syst. 106: 359–377 (1986)).
Distribution
A family of about 18 genera and 40–45 species in the tropics and subtropics.
Rhizomes
Rhizomes with or without scaly leaves Rhizomes with or without scaly leaves
Male
Male flowers: perianth segments (tepals) 3–9; stamens 2–12, free or united into a synandrium
Female
Female flowers: perianth absent or much reduced, 3–several-lobed and adnate to ovary
Gynoecium
Style 1, long, filiform, or absent Ovary inferior, 1–3-locular, ovules 1–3

Images

Balanophoraceae Rich. appears in other Kew resources:

First published in Mém. Mus. Hist. Nat. 8: 429. 1822 [18 Nov 1822] (1822)

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