1. Capparaceae Juss.

    1. This family is accepted.

[FTEA]

Capparidaceae, J. Elffers, R. A. Graham And G. P. Dewolf. Flora of Tropical East Africa. 1964

Habit
Herbs, shrubs or trees, sometimes scandent, glabrous, pubescent, glandular or scabrous, occasionally lepidote
Leaves
Leaves alternate or rarely opposite, simple or digitately 2–7-foliolate; leaflets entire, rarely serrate, dentate or panduriform
Stipules
Stipules present, minute or spiny, caducous or persistent
Inflorescences
Inflorescences terminal or axillary, racemose, often corymbose, or the flowers solitary or fascicled, often showy
Flowers
Flowers hermaphrodite or very rarely dioecious, regular or irregular
Calyx
Sepals 4–8, free or variously connate, valvate, imbricate or open in aestivation; receptacle cupular, funnel-shaped or cylindric, with an entire, undulate, dentate or fimbriate margin, sometimes very short
Corolla
Petals 4–16 or absent, sessile or clawed
Androecium
Stamens few to many, usually borne on a short or elongated androphore, all fertile or some without anthers; filaments free, equal or unequal, filiform; anthers 2-celled, oblong, longitudinally dehiscent
Gynoecium
Ovary sessile or more usually supported on a gynophore of varying length, 1- locular with parietal placentas or divided by spurious dissepiments into 2 or more locules; ovules few to many
Fruits
Fruit a capsule or berry, mostly oblong-cylindrical, globose or torulose, few- to many-seeded, rarely 1- seeded
Seeds
Seeds usually reniform or angular; endosperm none or scanty; embryo arcuate or incurved; cotyledons folded or convolute
Distribution
A medium-sized family of the tropics and subtropics, well represented in Africa and forming a conspicuous element of the dry-country flora
[FWTA]

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

Habit
Herbs, shrubs or trees, sometimes scandent
Leaves
Leaves alternate or rarely opposite, simple or digitately 3–7-foliolate; stipules when present minute or spiny
Flowers
Flowers mostly hermaphrodite, actinomorphic or rarely zygomorphic, hypogynous, axillary or terminal, variously arranged
Perianth
Perianth of calyx and corolla, or the latter absent
Calyx
Sepals free or partially united, imbricate or valvate, usually 4
Corolla
Petals 4 to many or absent
Nectaries
Torus elongated or short, rarely with an appendix
Androecium
Stamens few to many, sometimes some of them without anthers; filaments sometimes partially adnate to the torus; anthers 2-celled, longitudinally dehiscent
Gynoecium
Ovary sessile or more usually supported on a long or short gynophore, 1-celled with parietal placentas or divided into 2 or more cells by spurious dissepiments; ovules few to many
Fruits
Fruit a capsule or berry, sometimes the latter elongate or torulose
Seeds
Seeds usually reniform or angular; endosperm none or scanty; embryo arcuate or incurved
[FZ]

Capparidaceae, H. Wild. Flora Zambesiaca 1:1. 1960

Habit
Herbs, shrubs, trees or lianes
Leaves
Leaves alternate, simple or digitately 3––9-foliolate; stipules absent or rudimentary, rarely spiny
Inflorescences
Inflorescences terminal or axillary, of racemes, corymbs or panicles, or flowers solitary and axillary
Flowers
Flowers actinomorphic or zygomorphic, bisexual or unisexual by abortion, usually 4-merous, hypogynous
Receptacle
Receptacle cupular, funnel-shaped or cylindric with an entire, undulate, dentate or fimbriate margin, sometimes very short
Calyx
Sepals 3–4 (5)
Corolla
Petals 4 (0, 5, 6 or more)
Androecium
Stamens 5–? usually borne on a short or elongated androphore, sometimes accompanied by staminodes
Gynoecium
Style short or absent Ovary usually borne on a more or less elongated gynophore, usually 1-locular with 2 parietal placentas but sometimes 2-locular by the intrusion of the placentas or multi-locular; ovules 4–?
Fruits
Fruit a capsule or a berry
Seeds
Seeds reniform or subglobose, without endosperm; embryo usually curved
[NTK]

Cornejo, X. (2009). Neotropical Capparaceae.

Morphology
Description

Shrubs or trees, sometimes lianas, unarmed , glabrous or pubescent with simple , unicellular or uniseriate hairs, sometimes glandular , stellate , echinate or peltate - lepidote . Leaves usually evergreen , simple , rarely palmately 3-foliolate (Crateva L.), alternate , usually spiral or distichous , rarely opposite, entire , pinnately-veined, often coriaceous , subsessile to petiolate ; petioles lacking pulvini or pulvinate at one or both ends; stipules minute or lacking. Inflorescences racemose or corymbose, terminal and/or lateral , unbranched or often several compounded into terminal corymbose panicles, or flowers solitary in leaf axils; floral bracts present, highly reduced, deciduous . Flowers hypogynous, usually bisexual , sometimes cyclically unisexual, usually zygomorphic , rarely actinomorphic . Calyx aestivation open or closed, the sepals 4 (to 7 in Crateva), valvate , imbricate , decussate , with the lobes free , adnate, connate or totally fused, or the calyx bilabiate or spathaceous (Belencita H. Karst.). Corolla aestivation open or closed, the petals 4, imbricate or torsivae, rarely valvate (Calanthea (DC.) Miers), free , sessile or clawed, equal and cruciform. Stamens (4-)6-250(or more), as long as the petals to exserted above the petals, some sometimes shorter than the petals in dimorphic taxa, borne on a short to elongate androgynophore ; anthers usually basifixed, introrse, longitudinally dehiscent ; staminodia sometimes present. Hypanthium present or absent, the receptacle often more or less flat to conical, with 4 glands , scales or appendages, or with a nectariferous disk to bowl coating the hypanthium within. Ovary 2-4-carpellate, 1-4-locular with parietal placentation, or 2-8 "locular" with spurious septa from parietal placentas, superior , sessile or usually borne on a short to elongated gynophore ; style usually absent, the stigma 1, sessile , usually truncate and hemispherical to capitate (bilobed to undifferentiated only in Atamisquea Miers); ovules few to many, anatropous to campylotropous. Fruits subsessile to long-stipitate on a elongate gynophore , usually more or less fleshy , 1-locular, linear -cylindric to oblongoid or globose , dehiscent or indehiscent , capsular (2-4-valved), pepos, amphi-sarca or pseudoamphi-sarca; inner fruit wall ( endocarp ) often producing fleshy pulp surrounding arillate or the fiber-infiltratred, sarcotesta-covered seeds. Seeds 1 to usually many per fruit , usually cochleate to ± reniform , laterally somewhat compressed , with the testa invagination very short or shallow, barely if at all invaginated between the radicle and cotyledons, the testa hard, brittle, or very thin, the embryo white to cream, yellow or green, curved to almost straight, basically cochleate - reniform and incumbent, but usually with the cotyledons variously conduplicated and little to abundantly folded into each other and around the radicle; or the seeds subglobose and strongly anisocotylar, the major cotyledon massive, compact, specialized for store starches and the minor cotyledon reduced or even absent (Anisocapparis Cornejo & Iltis); endosperm obsolete, very thin.

Distribution
Distribution in the Neotropics
  • Southern United States (Texas and southern Florida) to northern Argentina, and the West Indies.
Diagnostic
Key differences from similar families

Capparaceae is related to Brassicaceae and Cleomaceae. Capparaceae differs from the two families by the:

  • Woodyhabit (vs. herbaceous).
  • Fleshy fruits, without replum (vs. dry fruits, with replum).
Notable genera and distinguishing features

Anisocapparis:

  • Plants glabrous or with simple trichomes throughout.
  • Recognized by the calyxdecussate, anisosepalous, the outer pair of sepals smaller.
  • Floral nectaries of 4 erect scales, strongly dimorphic (2 major and 2 minor).
  • Pollen grains finely reticulate.
  • Fruits pepos.
  • Seeds subglobose, the embryo highly anisocotylar, the major cotyledon subglobose, massive, compact, specialized for store starches, and the minor cotyledon rudimentary or absent.
  • Cataphylls present in the earlier stages of germination.

Atamisquea:

  • Plants with lepidote-peltate trichomes throughout.
  • Characterized by the branches bearing ± horizontally short straight and often spine -like branchlets.
  • Reduced leaf blades (0.5 to 3.5 cm long).
  • Calyx with sepals dimorphic, the outer pair valvate, wider and entirely enclosing the narrower and more delicate inner pair of sepals until or nearly to anthesis.
  • Style present, short.
  • Stigma bilobed, rather undifferentiated.
  • Fruits small (ca. 4-6 x 4-6 mm), apiculate at the apex; containing 1 to 2 small seeds.

Belencita:

  • Plants with stellate trichomes throughout.
  • Unique in the bialate or spathaceous calyx.
  • Fruits amphisarca.

Beautempsia:

  • Plants with stellate trichomes throughout.
  • Distinguished by the calyx hemisphaerical-cyathiform, with open aestivation, the lobes reduced.
  • Staminal nectaries present at the base of the filaments forming a nectariferous dish-shaped digitate pseudotorus.
  • Fruits pepos.

Calanthea:

  • Plants with stellate trichomes throughout.
  • Recognized by the sepals linear-ligulate to oblong, widely spaced, arranged on and covering the petal sutures, exposing the petals from very young buds.
  • Corolla with valvateaestivation.
  • Cotyledons 4-7 mm thick, slightly folded.
  • Fruits amphisarca.

Capparicordis:

  • Plants with stellate trichomes throughout.
  • Characterized by the leaves ovate to ovate-cordate, with (sub) palmatevenation at the base of blade.
  • Calyx 1-seriate, valvate.
  • Hypanthium present.
  • Petals yellow.
  • Stamens 4 to 8.
  • Nectary appendages linear to narrowly-triangular.
  • Fruits dehiscent, with the soft and very flexible fruit wall splitting and falling apart in 2 to 4 segments at maturity.

Capparidastrum:

  • Plants glabrous or with simple trichomes throughout
  • Recognized by the fleshy floral nectaries, usually rounded
  • Filaments of the stamens inserted on several alternate whorls on a hemispherical upper part of the androphore (seen at anthesis in fresh material)
  • This genus shares with Anisocapparis the simple hairs or glabrous pubescence, and the pepo fruits. However, Capparidastrum mainly differs from the latter by the calyx isosepalous, the valvate to imbricateaestivation, and the isocotylar seeds with convolute cotyledons

Colicodendron:

  • Plants with stellate trichomes throughout.
  • Calyx 1-seriate, valvate, ± cup-shaped, the sepals at the base fused forming a hypanthium, coated by a dentate or lobednectary dish, the sepal lobes free at the apex, developed, distinct and entirely enclosing the corolla in bud, their margins and tips touching each other from early bud until or nearly to anthesis.
  • Fruits amphisarca or pseudoamphisarca.

Crateva:

  • Plants glabrous or with furfuraceous trichomes throughout
  • Characterized by the leaves palmately 3-foliolate
  • Corolla with open aestivation
  • Floral nectary a dish- to bowl-shape coating the hypanthium within
  • Fruit amphisarca or pepos

Cynophalla:

  • Plants glabrous or with simple trichomes throughout.
  • Distinguished by the terminal branchlets with extra-floral axillary nectaries.
  • Usually distichous leaves.
  • Floral nectaries flat to somewhat concave, horizontally to widely divergent on the receptacle.
  • This genus shares with Anisocapparis the pubescence of simple hairs or glabrous and the decussate anisosepalous calyx type with the outer pair of sepals smaller. However, Cynophalla also differs from the latter genus by the pollen tectate-spinulose, and the capsular fruits, containing seeds with green, very convolute cotyledons.

Mesocapparis:

  • Lianas or scandent shrubs with stellate trichomes throughout.
  • This genus is characterized by the distichous leaves.
  • Solitary, axillary, ebracteate flowers.
  • Sepals imbricate in bud, the outer pair partially overlapping the inner pair, the inner pair exposed from very young bud.
  • Fruits pepos.

Monilicarpa:

  • Plants glabrous or with simple trichomes throughout.
  • Characterized by the bracts of-and/or near-the base of peduncles of inflorescences to 13 mm long, linear.
  • Floral nectaries cupular or disciform within a small hypanthium.
  • Petals fused on the edge or at the outer upper side of a cupular or disciform floral nectary.
  • Stamens with the filaments inserted in 2 alternate whorls on a slightly swollen androgynophore.
  • Pollen grains striate-reticulate.
  • Fruit capsular.
  • Monilicarpa additionally differs from Capparidastrum by the fruits moniliform or linear-torulose, essentially without pulp, the apices narrowly-attenuate, and the seeds arranged in one row and separated by the fruit -wall constrictions.

Morisonia:

  • Plants with stellate- to peltate-echinate trichomes throughout.
  • Recognized by the inflorescences cauliflorous and/or ramiflorous.
  • Sepals entirely fused, irregularly rupturing at anthesis into 2-3(-4) segments.
  • Fruits spherical amphisarca, the gynophores very thick and several times shorter than fruit body.

Neocalyptrocalyx:

  • Plants with variously stellate trichomes throughout.
  • Distinguished by a 2-seriate calyx with the outer pair of sepals valvate-connate, which encloses the inner pair until or nearly to anthesis, the sepal sutures usually not visible in flower buds.
  • Among the Neotropical Capparaceae, only Atamisquea has a somewhat similar 2-seriate calyx of closed aestivation with the outer pair of sepals valvate.
  • However, Neocalyptrocalyx differs from the latter by the stellate (vs. lepidote-peltate) trichomes, the more ascendent pattern of the branches, and the larger leaves.
  • The flower buds of Neocalyptrocalyx are circular in cross section and usually have the sepals smooth outside and the sutures not visible (vs. flower buds somewhat compressed laterally with the sepal sutures ± keeled, like an inverted U-shape arc on the buds in Atamisquea).
  • The flowers of Neocalyptrocalyx at anthesis have four nectary scales (vs. a dentatenectary cup in Atamisquea), the staminodes are absent (vs. present), and the ovaries have a sessilestigma, ± truncate at the apex (vs. ovaries at the apex and style sharply acute to subrostrate in Atamisquea).
  • Finally, the fruits of Neocalyptrocalyx are larger and many-seeded than Amphisarca, with a subwoody thick wall, 3-10 mm thick (vs. the small, and tardily dehiscent, pseudopeponoid fruits of Atamisquea).

Quadrella:

  • Plants with lepidote to variously stellate trichomes throughout.
  • This genus is distinguished by the calyx 1-seriate, valvate, dialisepalous.
  • Nectary scales arranged on a flat receptacle, the hypanthium absent.
  • Fruits capsular, usually linear-torulose to oblong, the inner fruit wall and pulp bright orange to bright-red.

Sarcotoxicum:

  • Plants with multiangulate-stellate, echinate to candelabroid trichomes throughout
  • Recognized by the imbricate sepals and the distinctive cucurbitoid amphisarca, containing a highly poisonous pulp
  • Due to the presence of stellate to candelabra trichomes, the calyces with 2-seriate sepals, and the seeds with white embryos of thinly convolute cotyledons, Sarcotoxicum is morphologically related to the also South American genera Mesocapparis and Neocalyptrocalyx
  • However, Sarcotoxicum differs from both by having flower buds cylindrically ellipsoid to cylindrically oblong-elliptic (vs. spherical to widely ovate-globose); corolla of imbricate (vs. siniestrorsely-torsivae) aestivation; flowers zygomorphic at anthesis, with the petals adaxially and laterally arranged, the stamens abaxially disposed, and the gynophore off-center (vs. flowers actinomorphic at anthesis, with the petals and stamens radially arranged, and the gynophore centered); and staminodia and a dentatenectary cup present (vs. staminodia and nectary cup absent, but 4 nectary scales present).

Steriphoma:

  • Shrubs or lianas with stellate trichomes throughout.
  • This genus is characterized by the calyx 1-seriate, tubular, subcylindric or funnelform, bright orange to orange-red (alive), the apical lobes 2-4, indistinct at the early bud stage.
  • Stamens (5-)6(-8).
  • Fruits indehiscent, decomposing at maturity, linear, ± torulose, oblong or cylindric to thickly bacciform.
Useful tips for generic identification
  • Pubescence type.
  • Presence or absence of extra-floral axillary nectaries.
  • Calyx aestivation.
  • Corolla aestivation.
  • Cotyledons type.
Other important characters
  • Stamens (in 95% of cases) numerous, usually exserted.
Distinguishing characters (always present)
  • Woody plants.
  • Leaves simple, or 3-foliolate in Crateva.
  • Floral bracts highly reduced, deciduous.
  • Flowers with 4 scales, glands or appendages; or a nectary dish/bowl within the hypanthium.
  • Ovary subtended by a usually exsertedgynophore.
  • Fruits without replum, usually fleshy.
General Description
General notes
  • Morphological studies and molecular data confirm that the American species traditionally identified as Capparis L. do not belong to that genus.
  • Species of this family are often dominant evergreen elements in dry forests.
  • The flowers are ephemeral, mostly nocturnal.
  • Some species (e.g. Quadrellacynophallophora (L.) Hutch., Sarcotoxicumsalicifolium (Griseb.) Cornejo & Iltis) are cultivated as ornamental.
  • The pulp of the fruits of several species of Capparidastrum and Neocalyptrocalyx is eaten by both humans and monkeys.
  • The pulp of the fruits of species of Calanthea and Sarcotoxicum is highly poisonous.
Number of genera

18 genera:

  • Anisocapparis Cornejo & Iltis
  • Atamisquea Miers
  • Beautempsia (Benth. & Hook.)Gaudich.
  • Belencita H. Karst.
  • Calanthea (DC.) Miers
  • Capparicordis Iltis & Cornejo
  • Capparidastrum (DC.) Hutch.
  • Colicodendron Mart.
  • Crateva L.
  • Cynophalla (DC.) J. Presl
  • Hispaniolanthus Cornejo & Iltis
  • Mesocapparis (Eichl.) Cornejo & Iltis
  • Monilicarpa Cornejo & Iltis
  • Morisonia L.
  • Neocalyptrocalyx Hutch.
  • Quadrella (DC. ) J. Presl
  • Sarcotoxicum Cornejo & Iltis
  • Steriphoma Spreng.
Status
  • Native.
Notes on delimitation
  • Capparaceae has been included in Brassicaceae s.l. (APG, 1998). Subsequent molecular studies (Hall et al., 2002, 2008) strongly support that Capparaceae s.s. must be considered a separate family.
Literature
Important literature

Candolle, A.P. De. 1824. Capparideae. Pp. 237-254 in: Prodromus systematis naturalis regni vegetabilis 1. Paris.

Cornejo, X. & H. H. Iltis. 2006. New combinations in Capparaceae sensu stricto for Flora of Ecuador. Harvard Pap. Bot. 11(1): 17--18.

Cornejo, X. & H. H. Iltis.2008a. Two new genera of Capparaceae: Sarcotoxicum and Mesocapparis stat. nov., and the reinstatement of Neocalyptrocalyx. Harvard Pap. Bot. 13(1): 103-116.

Cornejo, X. & H. H. Iltis. 2008b. New combinations in South American Capparaceae. Harvard Pap. Bot. 13(1): 117-120.

Cornejo, X. & H. H. Iltis. 2008c. A revision of the American species of the genus Crateva. (Capparaceae). Harvard Pap. Bot. 13(1): 121-135.

Cornejo, X. & H. H. Iltis. 2008d. Anisocapparis y Monilicarpa: dos nuevos géneros de Capparaceae de América del Sur. J. Bot. Res. Inst. Texas 2(1): 61-74.

Cornejo, X. & H. H. Iltis. 2008e. The reinstatement of Capparidastrum. Harvard Pap. Bot. 13(2): 229-236.

Cornejo, X. & H. H. Iltis. 2008f. A revision of Colicodendron Mart. (Capparaceae s.s.). J. Bot. Res. Inst. Texas, 2(1): 75-93.Cornejo, X. & H. H. Iltis. 2009. Hispaniolanthus: A new genus of Capparaceae endemic to Hispaniola. Harvard Pap. Bot. 14(1): 9-14.Cornejo, X. & H. H. Iltis. 2009. The reinstatement of Beautempsia (Capparaceae) and a key to the genera of Neotropical Capparaceae with variously stellate or peltate indumenta. J. Bot. Res. Inst. Texas 3(2): 683-689.Iltis, H.H. & X. Cornejo. 2010. Studies in the Capparaceae XXIX: Synopsis of Quadrella, a Mesoamerican a West Indies genus. Jour. Bot. Res. Inst. Texas 4(1): 117-132.EICHLER, A.W. 1865. Capparideae. Pages 237-292 in C.F.P. VON MARTIUS, ED., Flora brasiliensis 13(1). München.

HALL, J. C., K. J. SYTSMA, AND H. H. ILTIS. 2002. Phylogeny of Capparaceae and Brassicaceae based on chloroplast sequence data. Amer. J. Bot. 89: 1826--1842.

HALL, J. C., K. J. SYTSMA, AND H. H. ILTIS. 2008. Systematics of Capparaceae and Cleomaceae: an evaluation of the generic delimitations of Capparis and Cleome using plastid DNA sequence data. Botany 86: 682--696.

HUTCHINSON, J. 1967. Pages 303-317 in: The Genera of Flowering Plants (Angiospermae) Dicotyledones. Vol. 2. Clarendon Press, Oxford: 659 pp.

ILTIS, H. H. & X. CORNEJO. 2007a. Studies in the Capparaceae XXX. Capparicordis, a new genus from the Neotropics. Brittonia 59: 246--254.

Images

Capparaceae Juss. appears in other Kew resources:

First published in Gen. Pl. [Jussieu] 242. 1789 [4 Aug 1789] , as 'Capparides' (1789)

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 2018. 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