Basellaceae Raf.

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

Morphology General Habit

Subsucculent herbs with slender twining stems, glabrous

Morphology Leaves

Leaves alternate, entire or almost entire, usually petiolate, exstipulate

Morphology Reproductive morphology Inflorescences

Inflorescences of spikes, racemes or panicles, axillary or terminal

Morphology Reproductive morphology Flowers

Flowers small, actinomorphic, hermaphrodite or unisexual; bracts small; bracteoles 2–4, frequently appressed to base of perianth, sometimes winged

Morphology Reproductive morphology Flowers Perianth

Perianth 5-lobed, lobes united at base only or into a 5-lobed tube, imbricate, persistent in fruit

Morphology Reproductive morphology Flowers Androecium

Stamens 5, inserted opposite and at base of perianth lobes; filaments free; anthers dehiscing variously

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, unilocular; styles terminal, free or united or 3-fid; stigmas 3; ovule solitary, basal, short stalked

Morphology Reproductive morphology Fruits

Fruit indehiscent, enveloped by persistent frequently fleshy perianth

Morphology Reproductive morphology Seeds

Seeds solitary, globose; endosperm present

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

Perennial, ± fleshy to succulent, herbaceous or occasionally suffruticose vines, or sometimes ± erect plants

Morphology Roots

Roots fibrous to (at least sometimes) thickened

Morphology Stem

Stem usually glabrous, rarely asperous or when young puberulent, up to several metres long, sometimes producing subterranean or aerial tubers

Morphology Leaves

Leaves alternate simple, without stipules, petiolate; leaf blades entire, rarely dentate by glands

Morphology Reproductive morphology Inflorescences

Inflorescences axillary or terminal spikes, racemes, panicles, or dichasia

Morphology Reproductive morphology Inflorescences Bracts

Bracts thin, deciduous or persistent

Morphology Reproductive morphology Flowers Pedicel

Pedicels absent or present

Morphology Reproductive morphology Inflorescences Bracteoles

Bracteoles 2, opposite, at apex of pedicel or displaced due to elongation of pedicel, deciduous or persistent, usually thin, rarely lacking

Morphology Reproductive morphology Flowers

Flowers sessile to pedicellate, except calyx actinomorphic, bisexual or rarely functionally unisexual, chasmogamous or cleistogamous

Morphology Reproductive morphology Flowers Calyx

Sepals 2, valvate, opposite, alternating with bracteoles, petaloid, patent to erect, persistent and often somewhat accrescent in fruit, free from each other or partly connate, fused at least at base with petals, usually entire, occasionally with a gibba at base or a dorsal wing (in fruit), at anthesis greenish, whitish, yellowish, or reddish, in fruit pale, brownish, or ± black, membranous to rather thick, rarely fleshy

Morphology Reproductive morphology Flowers Corolla

Petals 5, imbricate, patent to erect, persistent and often somewhat accrescent in fruit, connate only at base or up to c. 2/3 of their length, usually entire, at anthesis greenish, whitish, yellowish, or reddish, in fruit pale, brownish, or ± black, membranous to rather thick, rarely fleshy

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens 5, epipetalous, connate and fused only at base or up to c. 3/4 of their length with petals; anthers dorsifixed or basifixed, tetrasporangiate, dithecal, extrorsely dehiscent by longitudinal slits or by short, apical slits; pollen spheroidal to cuboidal, perforate or rarely reticulate, spinulose or not, colpate or porate

Morphology Reproductive morphology Flowers Gynoecium Pistil

Pistil 1, 3-carpellate; ovary superior, syncarpous, unilocular with basal placentation; ovule 1, bitegmic, amphitropous to campylotropous; style 1, undivided to 3- parted almost to the base, or styles 3; stigmas linear to capitate or 3- lobed, rarely bifid or undifferentiated

Morphology Reproductive morphology Fruits

Fruit indehiscent, nut -like, usually smooth, rarely rugose, in cross section rounded to somewhat hexagonal, surrounded at base to completely enclosed by persistent perianth; embryo annular to cochleate .

Distribution

Anredera Juss.: Mexico to SE Brazil and Venezuela, and the Caribbean Islands. Basella L.: Central America to S Peru and E Brazil, and the Caribbean Islands (introduced). Tournonia Moq.: W Colombia to N Ecuador. Ullucus Caldas: Venezuela to N Argentina. All species of Anredera, Tournonia, and Ullucus are native, and all except a few widespread Anredera species are endemic; the Basella species found in the Neotropics is cultivated and naturalised.

Diagnostic

Distinguishing characters (always present): Small flowers with 2 petaloid sepals and 5 petals, subtended by 2 opposite bracteoles. Usually fleshy to succulent vines. Fruits nut-like and mostly enclosed by persistentperianth. Key to genera of Neotropical Basellaceae 1. Leaf margin dentate by glands; inflorescence a dichasium...Tournonia1. Leaf margin entire; inflorescence a raceme, spike, or panicle...2 2. Petals caudate at apex; anthers basifixed, dehiscent by short apical slits —Ullucus 2. Petals obtuse at apex; anthers dorsifixed, dehischent by longitudinal slits —3 3. Inflorescence axisfleshy; flowers sessile...Basella3. Inflorescence axis not fleshy; flowers pedicellate —Anredera

Note

Most species are found in open, dry habitats. Ullucus tuberosus Caldas is an important high Andean food crop, and Basellaalba L. is cultivated for its edible leaves. A few Anredera species are grown as ornamentals, also outside their native distributions. The homologies of the (1) bracteoles, (2) sepals, and (3) petals have been much debated without being definitely settled. They are sometimes referred to as (1) bracts, pedicellar bracts, lateral bracts, lower bractlets, or lower bracteoles; (2) upper bractlets, upper bracteoles, or involucral bracts; (3) sepals or tepals. Number of genera: 4 genera: Anredera (12 spp.). Basella (1 introduced sp.). Tournonia (1 sp.). Ullucus (1 sp.).

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

Morphology General Habit

Twiners with slender steins and rather fleshy alternate entire petiolate leaves; stipules absent

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, actinomorphic, in spikes, racemes or panicles; bracts small; bracteoles 2, often united to the base of the calyx

Morphology Reproductive morphology Flowers Calyx

Sepals 5, often coloured, almost free or united into a 5-lobed tube, imbricate, persistent in fruit

Morphology Reproductive morphology Flowers Corolla

Petals absent

Morphology Reproductive morphology Flowers Androecium

Stamens 5, inserted opposite and at the base of the sepals; filaments free, short; anthers with 2 parallel cells opening longitudinally

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, 1-celled, with a solitary basal shortly stalked ovule; style terminal, often deeply divided into 3 stigmas

Morphology Reproductive morphology Fruits

Fruit indehiscent, surrounded by the persistent often fleshy calyx

Morphology Reproductive morphology Seeds

Seed solitary, almost spherical, with a membranous testa; endosperm copious, surrounded by the spirally twisted embryo

Basellaceae, B. Verdcourt. Flora of Tropical East Africa. 1968

Morphology General Habit

Subsucculent, glabrous, twining herbs with slender stems and alternate, entire, petiolate or rarely sessile, exstipulate leaves

Morphology Reproductive morphology Flowers

Flowers regular, hermaphrodite or unisexual, in spikes, racemes or panicles; bracts small; bracteoles 2–4, often 2 adnate to the base of the perianth, sometimes wing-like

Morphology Reproductive morphology Flowers Perianth

Perianth 5-lobed; lobes imbricate, sometimes coloured, united into a tube below or almost free, persistent

Morphology Reproductive morphology Flowers Androecium

Stamens 5, opposite to the perianth-lobes, inserted at their base; filaments free, short; anthers versatile, variously dehiscing

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, 1-locular; ovule solitary, basal, shortly stalked, campylotropous; style terminal, simple or 3-fid, or 3 free styles

Morphology Reproductive morphology Fruits

Fruit indehiscent, surrounded by the persistent often fleshy perianth or winged bracteoles

Morphology Reproductive morphology Seeds

Seeds solitary, almost spherical; endosperm copious or almost absent, surrounded by the spirally twisted or semi-annular embryo

Eriksson, R. (2007). A Synopsis of Basellaceae. Kew Bulletin, 62(2), 297-320. Retrieved from http://www.jstor.org/stable/20443356

Type

Basella L.

Morphology General Habit

Perennial, ± fleshy to succulent, herbaceous or occasionally suffruticose vines, or sometimes ± erect plants

Morphology Roots

Roots fibrous to at least sometimes thickened

Morphology Stem

Stem usually glabrous, rarely asperous or when young puberulent, up to several metres long, sometimes producing subterranean or aerial tubers

Morphology Leaves

Leaves spirally arranged, simple, without stipules, petiolate; leaf blades entire, rarely dentate by glands

Morphology Reproductive morphology Inflorescences

Inflorescences axillary or terminal spikes, racemes, panicles, or dichasia

Morphology Reproductive morphology Inflorescences Bracts

Bracts thin, deciduous or persistent

Morphology Reproductive morphology Flowers Pedicel

Pedicels absent or present

Morphology Reproductive morphology Inflorescences Bracteoles

Bracteoles 2, opposite, at apex of pedicel or displaced due to elongation of pedicel, deciduous or persistent, usually thin, rarely lacking

Morphology Reproductive morphology Flowers

Flowers sessile to pedicellate, except calyx actinomorphic, bisexual or rarely functionally unisexual, chasmogamous or cleistogamous

Morphology Reproductive morphology Flowers Calyx

Sepals 2, valvate, opposite, alternating with bracteoles, petaloid, patent to erect, persistent and often somewhat accrescent in fruit, free from each other or partly connate, fused at least at base with petals, usually entire, occasionally with a gibba at base or a dorsal wing (in fruit), at anthesis greenish, whitish, yellowish, or reddish, in fruit pale, brownish, or ± black, membranotus to rather thick, rarely fleshy

Morphology Reproductive morphology Flowers Corolla

Petals 5, imbricate, patent to erect, persistent and often somewhat accrescent in fruit, connate only at base or up to c. 2/3 of their length, usually entire, at anthesis greenish, whitish, yellowish, or reddish, in fruit pale, brownish, or + black, membranous to rather thick, rarely fleshy

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens 5, epipetalous, connate and fused only at base or up to c. 3/4 of their length with petals; anthers dorsifixed or basifixed, tetrasporangiate, dithecal, extrorsely dehiscent by longitudinal slits or by short, apical slits; pollen spheroidal to cuboidal, perforate or rarely reticulate, spinulose or not, colpate or porate

Morphology Reproductive morphology Flowers Gynoecium Pistil

Pistil 1, 3-carpellate; ovary superior, syncarpous, unilocular with basal placentation; ovule 1, bitegmic, amphitropous to campylotropous; style 1, undivided to 3-parted almost to the base, or styles 3; stigmas linear to capitate or 3-lobed, rarely bifid or undifferentiated

Morphology Reproductive morphology Fruits

Fruit indehiscent, nut-like, usually smooth, rarely rugose, in cross section rounded to somewhat hexagonal, surrounded at base to completely enclosed by persistent perianth; embryo annular to cochleate.

Distribution

Basellaceae is a small pantropical and subtropical family comprising four genera and a total of 19 species. Three genera (Anredera, Tournonia, and Ullucus) are native to the New World, and the remaining one (Basella) to the Old World.

Ecology

Most species are succulent vines that prefer dry habitats.

Eriksson, R. (2009). Neotropical Basellaceae.

Morphology

Description

Perennial , ± fleshy to succulent , herbaceous or occasionally suffruticose vines, or sometimes ± erect plants. Roots fibrous to (at least sometimes) thickened. Stem usually glabrous , rarely asperous or when young puberulent , up to several metres long, sometimes producing subterranean or aerial tubers. Leaves alternate , simple , without stipules, petiolate ; leaf blades entire , rarely dentate by glands . Inflorescences axillary or terminal spikes, racemes, panicles, or dichasia. Bracts thin, deciduous or persistent . Pedicels absent or present. Bracteoles 2, opposite, at apex of pedicel or displaced due to elongation of pedicel , deciduous or persistent , usually thin, rarely lacking. Flowers sessile to pedicellate, except calyx actinomorphic , bisexual or rarely functionally unisexual, chasmogamous or cleistogamous. Sepals 2, valvate , opposite, alternating with bracteoles, petaloid , patent to erect , persistent and often somewhat accrescent in fruit , free from each other or partly connate , fused at least at base with petals, usually entire , occasionally with a gibba at base or a dorsal wing (in fruit ), at anthesis greenish, whitish, yellowish, or reddish, in fruit pale, brownish, or ± black, membranous to rather thick, rarely fleshy . Petals 5, imbricate , patent to erect , persistent and often somewhat accrescent in fruit , connate only at base or up to c. 2/3 of their length, usually entire , at anthesis greenish, whitish, yellowish, or reddish, in fruit pale, brownish, or ± black, membranous to rather thick, rarely fleshy . Stamens 5, epipetalous, connate and fused only at base or up to c. 3/4 of their length with petals; anthers dorsifixed or basifixed, tetrasporangiate, dithecal, extrorsely dehiscent by longitudinal slits or by short, apical slits; pollen spheroidal to cuboidal, perforate or rarely reticulate , spinulose or not, colpate or porate. Pistil 1, 3-carpellate; ovary superior , syncarpous, unilocular with basal placentation; ovule 1, bitegmic, amphitropous to campylotropous; style 1, undivided to 3- parted almost to the base, or styles 3; stigmas linear to capitate or 3- lobed , rarely bifid or undifferentiated. Fruit indehiscent , nut -like, usually smooth, rarely rugose , in cross section rounded to somewhat hexagonal, surrounded at base to completely enclosed by persistent perianth ; embryo annular to cochleate .

Distribution

Distribution in the Neotropics

• Anredera Juss.: Mexico to SE Brazil and Venezuela, and the Caribbean Islands.
• Basella L.: Central America to S Peru and E Brazil, and the Caribbean Islands (introduced).
• Tournonia Moq.: W Colombia to N Ecuador.
• Ullucus Caldas: Venezuela to N Argentina.

Diagnostic

Useful tips for generic identification

Key to genera of Neotropical Basellaceae

1. Leaf margin dentate by glands; inflorescence a dichasium...Tournonia
1. Leaf margin entire; inflorescence a raceme, spike, or panicle...2

2. Petals caudate at apex; anthers basifixed, dehiscent by short apical slits —Ullucus
2. Petals obtuse at apex; anthers dorsifixed, dehischent by longitudinal slits —3

3. Inflorescence axisfleshy; flowers sessile...Basella
3. Inflorescence axis not fleshy; flowers pedicellate —Anredera

Distinguishing characters (always present)

• Small flowers with 2 petaloid sepals and 5 petals, subtended by 2 opposite bracteoles.

Other important characters

• Usually fleshy to succulent vines.
• Fruits nut-like and mostly enclosed by persistentperianth.

General Description

Status

• All species of Anredera, Tournonia, and Ullucus are native, and all except a few widespread Anredera species are endemic; the Basella species found in the Neotropics is cultivated and naturalised.

General notes

• Most species are found in open, dry habitats. Ullucus tuberosus Caldas is an important high Andean food crop, and Basellaalba L. is cultivated for its edible leaves. A few Anredera species are grown as ornamentals, also outside their native distributions.
• The homologies of the (1) bracteoles, (2) sepals, and (3) petals have been much debated without being definitely settled. They are sometimes referred to as (1) bracts, pedicellar bracts, lateral bracts, lower bractlets, or lower bracteoles; (2) upper bractlets, upper bracteoles, or involucral bracts; (3) sepals or tepals.

Number of genera

4 genera:

• Anredera (12 spp.).
• Basella (1 introduced sp.).
• Tournonia (1 sp.).
• Ullucus (1 sp.).

Literature

Important literature

Eriksson, R. 2004. Basellaceae. In: N. Smith et al. (eds), Flowering plants of the Neotropics, pp. 44-45, Princeton University Press, Princeton.

Eriksson, R. 2007. A synopsis of Basellaceae. Kew Bulletin 62: 297-320.

Sperling, C. R. 1987. Systematics of the Basellaceae. Unpublished Ph.D. dissertation, Harvard University, Cambridge.

Sperling, C. R. & Bittrich, V. 1993. Basellaceae. In: K. Kubitzki et al. (eds), The families and genera of vascular plants 2: 143-146, Springer-Verlag, Berlin.