Combretaceae

Taxonomy

Family Name: Combretaceae R. Brown

Synonym(s): Bucidaceae Spreng.; Myrobalanaceae Martinov; Strephonemataceae Venkat. & Prak. Rao, nom. inval.; Terminaliaceae J. St.-Hil.

Common Name(s): amaranth family

*Number of genera/species: 10/530

List of genera records in GRIN-Global

Disseminule

fruit

Description

Fruit a drupedrupe:
(indehiscent drupe) a fleshy, indehiscent fruit with one more hard pits enclosing seeds, derived from single, superior, simple or compound ovary; (dehiscent drupe) a fruit with a dry or fibrous to fleshy or leathery outer husk that early to tardily breaks apart (or opens), exposing one or more nutlike pits enclosing the seeds
, acheneachene:
a dry, indehiscent, one-seeded fruit, with seed attached to pericarp at a single point, derived from a single, superior, simple or compound, one-loculed ovary
, or samarasamara:
a winged, indehiscent, dry fruit containing a single (rarely two) seed(s)
, (technically a pseudocarp as the outer layer is derived from the hypanthium, except in Strephonema, which is derived from a semi-inferior ovary and only the base is hypanthial tissue), 5–100 mm, rarely to 130 mm, globoseglobose:
3D shape—more or less spherical
to rhomboid, tereteterete:
approximately circular in cross section; width and thickness approximately equal
, star-shapedstar-shaped:
(of fruit), having a distal aspect like the stylized shape of a star, usually found in aggregate versus simple fruit
, rectangular, flattened, or compressedcompressed:
flattened; in grasses, used to denote compression (not necessarily flattened) either laterally or dorsiventrally
on one-side in transection, often beakedbeak:
a usually firm, terminal appendage, sometimes tapered
, usually 2–5 winged, longitudinally or rarely encircling seed body, if not winged then often ribbedribbed:
surface relief—wide, prominent, linear ridges that are generally rounded and longitudinally situated on the surface
or ridgedridged:
surface relief—raised, thick ridges, sharp edged or rounded, usually in a series that may cover the entire surface
, with one seed, often  calyxcalyx:
the outer whorl of the perianth; all the sepals of a flower
and/or stylestyle:
in a flower, the narrow and elongated part of the pistil between the stigma and the ovary; sometimes persisting in fruit
persistent. Pericarppericarp:
fruit wall or fruit coat
variously colored, dulldull:
reflecting only a low proportion of incident light, with no apparent sheen
or shinyshiny:
uniformly reflecting a high proportion of incident light at all angles
, woodywoody:
texture—consisting mainly of indurate lignified tissues, characteristic of or resembling wood
, corkycorky:
firm, relatively light, discontinuous but strongly cohesive, and resilient
, leatheryleathery:
texture—moderately thick, tough, and very pliable
, paperypapery:
texture—papyraceous, chartaceous; very thin, pliable, and readily torn; like paper
or fleshy, glabrousglabrous:
without hairs
or pubescentpubescent:
surface relief—bearing hairs
, often with white, yellow or red scales, sometimes wrinkledwrinkled:
surface relief—shallow, irregular folds and furrows covering the surface; appearing overall though crumpled and then spread out
, striatestriate:
surface relief—having fine, parallel lines, grooves or ridges
, or veiny, rarely with hornshorn:
a straight or curved, slenderly conic or conoidal protrusion, resembling an animal horn
or tubercles.

In several genera, wings are derived from accrescentaccrescent:
growing continuously
calyces, Calycopteris, or from prophylls, Dansiea and Macropteranthes. In Conocarpus, drupesdrupe:
(indehiscent drupe) a fleshy, indehiscent fruit with one more hard pits enclosing seeds, derived from single, superior, simple or compound ovary; (dehiscent drupe) a fruit with a dry or fibrous to fleshy or leathery outer husk that early to tardily breaks apart (or opens), exposing one or more nutlike pits enclosing the seeds
or nutletsnutlet:
˜achene
densely clustered into a head.

Seeds globoseglobose:
3D shape—more or less spherical
, fusiformfusiform:
spindle-shaped; broadest at the middle and tapering at both ends
, or angularangular:
2D shape—having sides that meet at acute or obtuse angles  
, 2–40 mm long. Seed coat brown, membranousmembranous:
texture—extremely thin, pliable, and fairly tough
, glabrousglabrous:
without hairs
, ribbedribbed:
surface relief—wide, prominent, linear ridges that are generally rounded and longitudinally situated on the surface
, lobed, striatestriate:
surface relief—having fine, parallel lines, grooves or ridges
, pittedpitted:
surface relief—surface with small depressions in which the areas between the hollows do not take on the appearance of a true reticular net
, reticulatereticulate:
surface relief—netted, raised walls or concave grooves forming a net-like surface pattern with flat, concave, or convex interspaces
, or wrinkledwrinkled:
surface relief—shallow, irregular folds and furrows covering the surface; appearing overall though crumpled and then spread out
.

Embryo well developed, completely filling seed cavity, straight, sometimes green. Cotyledons usually 2 but if fused, may appear as 1 (Combretum spp.), or 3–5 in Terminalia spp. Cotyledons usually variously folded or spirally twisted, sometimes not folded, rarely massive and conduplicate (Strephonema, Combretum spp.).

Endosperm absent.

Identification features

Fruit
Type drupedrupe:
(indehiscent drupe) a fleshy, indehiscent fruit with one more hard pits enclosing seeds, derived from single, superior, simple or compound ovary; (dehiscent drupe) a fruit with a dry or fibrous to fleshy or leathery outer husk that early to tardily breaks apart (or opens), exposing one or more nutlike pits enclosing the seeds
, acheneachene:
a dry, indehiscent, one-seeded fruit, with seed attached to pericarp at a single point, derived from a single, superior, simple or compound, one-loculed ovary
, samarasamara:
a winged, indehiscent, dry fruit containing a single (rarely two) seed(s)
Size range 5–100 mm long, rarely to 130 mm
Shape(s) globoseglobose:
3D shape—more or less spherical
, ellipsoidellipsoid:
3D shape—elliptic
, fusiformfusiform:
spindle-shaped; broadest at the middle and tapering at both ends
, ovoidovoid:
3D shape—ovate
, obovoidobovoid:
3D shape—obovate
, cylindricalcylindrical:
3D shape—a cylinder, with parallel sides and a circular cross-section; tubular or rod-shaped
, pear-shaped, rhomboid, sometimes short stipitatestipitate:
borne on a stalk
Texture woodywoody:
texture—consisting mainly of indurate lignified tissues, characteristic of or resembling wood
, corkycorky:
firm, relatively light, discontinuous but strongly cohesive, and resilient
, leatheryleathery:
texture—moderately thick, tough, and very pliable
, paperypapery:
texture—papyraceous, chartaceous; very thin, pliable, and readily torn; like paper
, fleshy
Surface relief usually 2–5 winged, ribbedribbed:
surface relief—wide, prominent, linear ridges that are generally rounded and longitudinally situated on the surface
or lobed, glabrousglabrous:
without hairs
or pubescentpubescent:
surface relief—bearing hairs
, sometimes with white, yellow, or red scales (lepidotelepidote:
bearing scales, scaly, squamate, squamose
)
Color(s) black, purple, blue, green, brown, yellow, vivid pink or pink, red
Unique features Usually winged or ribbedribbed:
surface relief—wide, prominent, linear ridges that are generally rounded and longitudinally situated on the surface
drupesdrupe:
(indehiscent drupe) a fleshy, indehiscent fruit with one more hard pits enclosing seeds, derived from single, superior, simple or compound ovary; (dehiscent drupe) a fruit with a dry or fibrous to fleshy or leathery outer husk that early to tardily breaks apart (or opens), exposing one or more nutlike pits enclosing the seeds
, achenesachene:
a dry, indehiscent, one-seeded fruit, with seed attached to pericarp at a single point, derived from a single, superior, simple or compound, one-loculed ovary
, or samarassamara:
a winged, indehiscent, dry fruit containing a single (rarely two) seed(s)
often with persistent calyxcalyx:
the outer whorl of the perianth; all the sepals of a flower
and/or stylestyle:
in a flower, the narrow and elongated part of the pistil between the stigma and the ovary; sometimes persisting in fruit
at apexapex:
the point farthest from the point of attachment, or the "tip" of an organ
enclosing a solitary seed completely filledfilled:
(of embryo) embryo fills the seed interior with the food reserves stored in the cotyledons; small amounts of endosperm may remain around the embryo
by the embryo..
Seed
Size range 2–40 mm long
Shape(s) globoseglobose:
3D shape—more or less spherical
, fusiformfusiform:
spindle-shaped; broadest at the middle and tapering at both ends
, angularangular:
2D shape—having sides that meet at acute or obtuse angles  
Surface relief smooth, ribbedribbed:
surface relief—wide, prominent, linear ridges that are generally rounded and longitudinally situated on the surface
, ridgedridged:
surface relief—raised, thick ridges, sharp edged or rounded, usually in a series that may cover the entire surface
Color(s) brown
Unique features Seeds, rarely disseminulesdisseminule:
detachable plant part capable of being disseminated and of propagating, commonly a seed or fruit
, without endosperm, and seed cavity completely filledfilled:
(of embryo) embryo fills the seed interior with the food reserves stored in the cotyledons; small amounts of endosperm may remain around the embryo
with the embryo.
Other
Embryo well developed, completely filling seed cavity, straight, sometimes green. Cotyledons usually 2 but if fused, may appear as 1 (Combretum spp.), or 3–5 in Terminalia spp. Cotyledons usually variously folded or spirally twisted, sometimes not folded, rarely massive and conduplicate (Combretum spp., Strephonema). 
Nutritive tissue endosperm absent

Distribution

Pantropical

Distribution map courtesy of Angiosperm Phylogeny Website.

References

Davidse et al. 2009–2018Davidse et al. 2009–2018:
Davidse GM, Sousa Sánchez M, Knapp S. and Chiang Cabrera F, eds. 2009–2018. Flora Mesoamericana. Missouri Botanical Garden, St. Louis, MO. Accessed: January–April 2024. URL: http://legacy.tropicos.org/Project/FM
; Flora of Australia 2021+Flora of Australia 2021+:
Flora of Australia. Australian Biological Resources Study, Canberra. Accessed January 2021–March 2024. URL: http://www.ausflora.org.au
; Flora of North America Editorial Committee 1993+Flora of North America Editorial Committee 1993+:
Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico [Online]. 22+ vols. Flora of North America Association, New York and Oxford. Accessed January-March 2024. URL: http://beta.floranorthamerica.org.
; Hutchinson and Dalziel 1954–1972Hutchinson and Dalziel 1954–1972:
Hutchinson J and Dalziel JM, revised by Keay RWJ and Hepper FN. 1954–1972. Flora of West Tropical Africa, ed. 2, 3 Vols. 2300 pp.
; Liben 1968Liben 1968:
Liben L. 1968. Combretaceae. In: Flore du Congo, du Rwanda et du Burundi, Spermatophytes. Bruxelles, Jardin botanique national de Belgique. [version PDF, 10.5281/zenodo.4514585]
Kirkbride et al. 2006Kirkbride et al. 2006:
Kirkbride JH, Jr, Gunn CR, and Dallwitz MJ. 2006. Family guide for fruits and seeds, vers. 1.0. Accessed September 2020-January 2022. URL: https://nt.ars-grin.gov/seedsfruits/keys/frsdfam/index.cfm .
; Kubitzki et al. 1990+Kubitzki et al. 1990+:
Kubitzki K et al., eds. 1990+. The families and genera of vascular plants. 7+ vols. Berlin etc.
; Takhtajan 2009Takhtajan 2009:
Takhtajan A. 2009. Flowering plants: Second edition. Springer Nature, Switzerland. 871 pp.
; Tomlinson 2016Tomlinson 2016:
Tomlinson PB. 2016. The Botany of Mangroves. 2nd ed. Cambridge University Press, Cambridge. Family: Combretaceae, pp. 228-241.
; Zhengyi et al. 2004+Zhengyi et al. 2004+:
Zhengyi W, Raven PH, and Deyuan H. 2004+. Flora of China [online]. 25 vols. Science Press, Beijing China & Missouri Botanical Garden, St. Louis USA. Accessed January–March 2024. http://flora.huh.harvard.edu/china/

*The number of genera and species is based on Christenhusz and Byng 2016Christenhusz and Byng 2016:
Christenhusz MJM and Byng JW. 2016. The number of known plant species in the world and its annual increase. Phytotaxa 261 (3): 201-217. https://doi.org/10.11646/phytotaxa.261.3.1
, which may differ from the number of genera in GRIN-Global.

  Fruits:   Combretum  sp.; Photo by A. Margina, USDA APHIS PPQ, imageID.idtools.org
Fruits: Combretum sp.; Photo by A. Margina, USDA APHIS PPQ, imageID.idtools.org
  Fruits:   Combretum  sp.; Photo by N. Diaz, USDA APHIS PPQ, imageID.idtools.org
Fruits: Combretum sp.; Photo by N. Diaz, USDA APHIS PPQ, imageID.idtools.org
  Embryo:   Combretum micranthum ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Combretum micranthum; Illustration by K. Parker, Kirkbride et al. (2006)
  Fruits:   Conocarpus erectus ; Photo by J. St. John, commons.wikimedia.org
Fruits: Conocarpus erectus; Photo by J. St. John, commons.wikimedia.org
  Fruit, seed:   Laguncularia racemosa ; Photo by K.E. Clancy, USDA APHIS PPQ, imageID.idtools.org
Fruit, seed: Laguncularia racemosa; Photo by K.E. Clancy, USDA APHIS PPQ, imageID.idtools.org
  Embryo:   Laguncularia racemosa ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Laguncularia racemosa; Illustration by K. Parker, Kirkbride et al. (2006)
  Fruit:     Macropteranthes leichhardtii ; Photo by M. Fagg, Atlas of Living Australia, ala.org.au
Fruit: Macropteranthes leichhardtii; Photo by M. Fagg, Atlas of Living Australia, ala.org.au
  Fruits:   Terminalia   amazonia ; Photo by A. Margina, USDA APHIS PPQ, imageID.idtools.org
Fruits: Terminalia amazonia; Photo by A. Margina, USDA APHIS PPQ, imageID.idtools.org
  Fruits:   Terminalia amazonia ; Photo by N. Diaz, USDA APHIS PPQ, imageID.idtools.org
Fruits: Terminalia amazonia; Photo by N. Diaz, USDA APHIS PPQ, imageID.idtools.org
  Fruits:   Terminalia canescens ; Photo by J.W. Wrigley, Atlas of Living Australia, ala.org.au
Fruits: Terminalia canescens; Photo by J.W. Wrigley, Atlas of Living Australia, ala.org.au
  Fruit:   Terminalia catappa ; Photo by M. Fagg, Atlas of Living Australia, ala.org.au
Fruit: Terminalia catappa; Photo by M. Fagg, Atlas of Living Australia, ala.org.au
  Fruit, seed:   Terminalia catappa ; Photo by S. Hurst, USDA-NRCS Plants Database
Fruit, seed: Terminalia catappa; Photo by S. Hurst, USDA-NRCS Plants Database
  Fruit:   Terminalia chebula , dried drupe; Photo by K.E. Clancy, USDA APHIS PPQ, imageID.idtools.org
Fruit: Terminalia chebula, dried drupe; Photo by K.E. Clancy, USDA APHIS PPQ, imageID.idtools.org
  Fruits:   Terminalia   chebula ; Photo by K.E. Clancy, USDA APHIS PPQ, imageID.idtools.org
Fruits: Terminalia chebula; Photo by K.E. Clancy, USDA APHIS PPQ, imageID.idtools.org
  Fruit:   Terminalia melanocarpa ; Photo by M. Fagg, Atlas of Living Australia, ala.org.au
Fruit: Terminalia melanocarpa; Photo by M. Fagg, Atlas of Living Australia, ala.org.au
  Embryo:   Terminalia oblongata;  Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Terminalia oblongata; Illustration by K. Parker, Kirkbride et al. (2006)
  Fruits:   Terminalia   paniculata ; Photo by C. Southwick, Dried Botanical ID
Fruits: Terminalia paniculata; Photo by C. Southwick, Dried Botanical ID
  Fruits:   Terminalia porphyrocarpa ; Photo by M. Fagg, Atlas of Living Australia, ala.org.au
Fruits: Terminalia porphyrocarpa; Photo by M. Fagg, Atlas of Living Australia, ala.org.au
  Fruits:   Combretum indicum ; Photo by C. Southwick, Dried Botanical ID
Fruits: Combretum indicum; Photo by C. Southwick, Dried Botanical ID
  Embryo:   Quisqualis indica ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Quisqualis indica; Illustration by K. Parker, Kirkbride et al. (2006)