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A SYSTEMATIC REVISION OF Cristina Inocencio, Diego Rivera, 3
M Concepcio ´nObo ´n, Francisco Alcaraz, 3
a
4
CAPPARIS SECTION CAPPARIS
and Jose-Antonio Barren ˜a 3
(CAPPARACEAE) 1, 2
ABSTRACT
A systematic revision of Capparis sect. Capparis, from western and Central Asia, North Africa, and Europe, is presented
here. The taxonomy of this section has been approached combining morphological, biogeographical and molecular data when
available. Ten species are recognized, including two new species, Capparis atlantica and C. zoharyi. In addition, four new
subspecies are presented: Capparis ovata subsp. myrtifolia, C. parviflora subsp. sphaerocarpa, C. sicula subsp. mesopotamica,
and C. sicula subsp. sindiana. Lectotypes are designated for C. aegyptia, C. hereroensis, C. mucronifolia, C. elliptica, C.
mucronifolia Boiss. subsp. rosanoviana, C. rupestris, C. ovata, C. parviflora, C. spinosa var. canescens, C. sicula subsp.
herbacea, and C. sicula subsp. leucophylla. A full taxonomic treatment, keys, and distribution maps of the recognized species
are provided. The two new species are illustrated.
Key words: Capparaceae, Capparis, taxonomy.

Capparis (Kappariz) is a name coined by Theo- transferred to section Monostichocalyx Radlk. and
phrastus (4th century BC) and endorsed by Dioscor- section Busbeckea (Endl.) Benth. & Hook. Subsection
ides (1st century AD). It seems to have come into wide Seriales DC. Racemose inflorescences or in series on
use after the spread of the Arab culture in the Middle the stems. Representative species: C. zeylanica L., C.
Ages. The Genus Capparis was created by Linnaeus acuminata Willd., C. quadriflora DC., C. rotundifolia
(1753, 1754) with the description of Capparis spinosa Rottl., C. brevispina DC. Subsection Corymbosae DC.
L. and other Capparis species (Jarvis et al., 1993). Corymbose inflorescences. Representative species: C.
Capparis comprises around 250 species distributed in sepiaria L., C. umbellata Brown ex DC., C. incanescens
tropical and subtropical zones of southern America, DC. subsection Octandrae DC. Flowers with 8
Europe, Africa, Madagascar, Asia, Australia, and the stamens. Representative species: C. racemulosa DC.,
Pacific Islands (Willis, 1988). C. oleoides Burch., C. coriacea Burch.
New World taxa are included in five sections.
Section 2. Capparidastrum DC. Stems generally
TAXONOMIC HISTORY
thorny, oval sepals without gland, and short gyno-
Candolle (1824), who proposed a sectional division phore. Representative species: C. brasiliana DC., C.
of the genus, provided the first comprehensive macrophylla HBK, C. cuneata DC. Section 3.
systematic approach to Capparis. Old World taxa Cynophalla DC. Stems generally thorny, oval sepals
belong to Section 1. Capparis (5 Eucapparis DC.). possessing basal gland or foveola, and large gyno-
Representative species: C. spinosa L. with four phore. Representative species: C. cynophallophora L.,
subsections. Subsection Capparis (5 Pedicellares C. guayaquilensis HBK, C. amplissima Lam. Section
DC.). Flowers always solitary at leaf axils. In addition 4. Calanthea DC. Rounded fruit section, thin sepals,
to C. spinosa and related species, other representa- linear, and sharp. Representative species: C. nemo-
tives include species such as C. cartilaginea Decne. rosa Jacq., C. pulcherrima Jacq. Section 5. Breynias-
Some species with geminate flowers or in bunches of trum DC. Triangular sepals, sharp; always very short
three, rarely alone, such as C. horrida L. f., C. gynophore. Representative species: C. ferruginea L.,
pubiflora DC., and C. canescens DC., were later C. incana HBK, C. indica (L.) Rawc. & Rendle.

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This work was part of the Ph.D. Dissertation presented by C. Inocencio to the Faculty of Biology, Murcia, Spain. We thank
the following herbaria and their staff for providing loans, access to collections, and assistance: BISH, BM, BR, C, E, G, HUB,
HUJ, K, JE, JEPS, LIV, MA, MARSSJ, MUB, OXF, P, RNG, RSA, UMH, and US. We thank Pedro Perales for drawing
Figure 1. Special gratitude is due to Jochen Mu ¨ller (JE), Vladimir Dorofeev (LE), Serena Marner (OXF), Donna Young (LIV),
and Aljos Farjon (K) for their help in finding type material. This research was supported by a grant from the Spanish Ministry
of Education and Culture and by the project AGF96-1040 of DGICYT.
2
The editors of the Annals thank Sophia Balcomb for her editorial contribution to this article.
3
Departamento de Biologı´a Vegetal, Facultad de Biologı´a, Universidad de Murcia, E-30100 Espinardo, Murcia, Spain.
drivera@um.es.
4 Departamento de Biologı´a Aplicada, Escuela Polite ´cnica Superior de Orihuela, Universidad Miguel Herna ´ndez, E-03312
Orihuela (Alicante), Spain.
ANN.MISSOURI BOT.GARD. 93: 122–149. PUBLISHED ON 31 MAY 2006.