Neotropical Capparaceae

Xavier Cornejo

New York Botanical Garden, Bronx, New York, USA. 

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.

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.

Distribution in the Neotropics

  • Southern United States (Texas and southern Florida) to northern Argentina, and the West Indies.

Distinguishing characters (always present)

Other important characters

  • Stamens (in 95% of cases) numerous, usually exserted.

Key differences from similar families

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

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.

Useful tips for generic identification

Notable genera and distinguishing features

Anisocapparis:

  • Plants glabrous or with simple trichomes throughout.
  • Recognized by the calyx decussate, 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 valvate aestivation.
  • Cotyledons 4-7 mm thick, slightly folded.
  • Fruits amphisarca.

Capparicordis:

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 imbricate aestivation, 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 lobed nectary 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 dentate nectary cup in Atamisquea), the staminodes are absent (vs. present), and the ovaries have a sessile stigma, ± 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:

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 dentate nectary 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.

Status

  • Native.

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. Quadrella cynophallophora (L.) Hutch., Sarcotoxicum salicifolium (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.

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.

How to cite

Cornejo, X. (2009). Neotropical Capparaceae. In: Milliken, W., Klitgård, B. & Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://www.kew.org/science/tropamerica/neotropikey/families/Capparaceae.htm.

Click images to enlarge


Fruits and flower buds of Beautempsia avicennifolia © Xavier Cornejo, New York Botanical Garden.



Inflorescence of Capparicordis crotonoides © Xavier Cornejo, New York Botanical Garden.



Leafy branch and inflorescences of Capparicoris crotonoides © Xavier Cornejo, New York Botanical Garden.



Sectioned mature fruit of Capparidastrum discolor © Xavier Cornejo, New York Botanical Garden.



Sectioned mature fruit of Capparidastrum frondosum © Xavier Cornejo, New York Botanical Garden.



Flower of Capparidastrum frondosum © Xavier Cornejo, New York Botanical Garden.



Infructescence of Capparidastrum frondosum © Xavier Cornejo, New York Botanical Garden.



Inflorescence of Colicodendron scabridum © Xavier Cornejo, New York Botanical Garden.



Sectioned mature fruit of Colicodendron scabridum © Xavier Cornejo, New York Botanical Garden.



Axillary nectary glands of Cynophalla amplissima © Xavier Cornejo, New York Botanical Garden.



Flowerbuds of Cynophalla amplissima © Xavier Cornejo, New York Botanical Garden.



Flowered branch of Cynophalla amplissima © Xavier Cornejo, New York Botanical Garden.



Sectioned mature fruit of Cynophalla heterophylla displaying the purple pulp and seeds surrounded by white arils and containing green cotyledons © Xavier Cornejo, New York Botanical Garden.



Terminal branchlet of Cynophalla mollis bearing leaves and mature fruits © Xavier Cornejo, New York Botanical Garden.



Inflorescence of Cynophalla sclerophylla © Xavier Cornejo, New York Botanical Garden.