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The calybium and the cupule make up the
accessory fruit An accessory fruit is a fruit in which some of the flesh is derived not from the floral ovary but from some adjacent tissue exterior to the carpel.Esau, K. 1977. ''Anatomy of seed plants''. John Wiley and Sons, New York. Accessory fruits are u ...
of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s in the family Fagaceae. These two parts derive from different flower components. The cupule holds and protects the
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
during its growth and maturation. In some genera (e.g. '' Lithocarpus,
Quercus An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably ...
''), it only partly encloses the single nut, while in others (e.g. '' Castanea,
Fagus Beech (''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to temperate Europe, Asia, and North America. Recent classifications recognize 10 to 13 species in two distinct subgenera, ''Engleriana'' and ''Fagus''. The ''Engle ...
''), it fully encloses the two or more nuts, and splits open at maturity into four valves to release the nuts. It is derived from the vegetative part of the flower (its attachment to the rest of the plant). It is covered by numerous scales. In some (e.g. ''Castanea''), the scales are developed into sharp spines, giving the nut protection from squirrels and other seed predators, while in others (e.g. most ''Quercus''), they are not. In ''Lithocarpus'', the cupule is very hard and bone-like in texture. The calybium (plural: calybia) is the fruit proper. It develops from an
inferior ovary In the flowering plants, an ovary is a part of the female reproductive organ of the flower or gynoecium. Specifically, it is the part of the pistil which holds the ovule(s) and is located above or below or at the point of connection with the ba ...
, meaning it is initially encased in the future cupule. Technically the calybium is a
nut Nut often refers to: * Nut (fruit), fruit composed of a hard shell and a seed, or a collective noun for dry and edible fruits or seeds * Nut (hardware), fastener used with a bolt Nut or Nuts may also refer to: Arts, entertainment, and media Co ...
, as its ovary wall becomes dry with the embryo loosely enclosed inside, and remains closed until germination. In the related family
Betulaceae Betulaceae, the birch family, includes six genera of deciduous nut-bearing trees and shrubs, including the birches, alders, hazels, hornbeams, hazel-hornbeam, and hop-hornbeams numbering a total of 167 species. They are mostly natives of the ...
, notably in the genera ''
Carpinus Hornbeams are hardwood trees in the flowering plant genus ''Carpinus'' in the birch family Betulaceae. The 30–40 species occur across much of the temperate regions of the Northern Hemisphere. Origin of names The common English name ''hornbea ...
'' and ''
Corylus The hazel (''Corylus'') is a genus of deciduous trees and large shrubs native to the temperate Northern Hemisphere. The genus is usually placed in the birch family Betulaceae,Germplasmgobills Information Network''Corylus''Rushforth, K. (1999). ...
'', the cupule is replaced by an
involucre In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or o ...
, which differs in being more leafy in appearance, but performs a similar role in protecting the developing nuts.


References

* (2003): A Miocene rodent nut cache in coastal dunes of the Lower Rhine Embayment, Germany. '' Palaeontology'' 46(6): 1133-1149. {{doi, 10.1046/j.0031-0239.2003.00337.x Fagaceae Fruit morphology