REVIEW OF FOOD AND MEDICINAL USES OF CAPPARIS L. SUBGENUS CAPPARIS (CAPPARIDACEAE) 1

DIEGO RIVERA, CRISTINA INOCENCIO, CONCEPCION OBON, AND

FRANCISCO ALCARAZ

Rivera, D., Inocencio, C., Alcaraz, F. (Department of Plant Biology, University of Murcia, 30100 Espinardo, Murcia, Spain. e-mail: drivera@um.es) Ob6n, C. (Department of Applied Biology, EPSO, University Miguel Herndndez, 03312 Orihuela, Alicante, Spain). REVIEW OF FOOD AND MEDICINAL USES OF CAPPARIS L. SUBGENUS CAPPARIS (CAPPARIDACEAE). Economic Botany 57(4):515-534, 2003. Capers of commerce are immature flower buds which have been pickled either in vinegar or preserved in granular salt. Semi-mature fruits and young shoots with small leaves may also be pickled for use as a condiment. The use of capers can be traced to the prehistory. Although Capparis spinosa from the western Mediterranean is the most widely used species, the subgenus comprises 23 species and subspecies occupying large territories from the Atlantic coasts to the Pacific in the Old World. We have recorded medicinal and food uses for 19 species.

REVISION DE LOS Usos ALIMENTARIOS Y MEDICINALES DE CAPPARIS SUBGF.NERO CAPPARIS (CAP- PARIDACEAE). I_ZIS alcaparras del comercio son botones florales encurtidos en vinagre o pres- ervados en sal gruesa. Los frutos inmaduros y los brotes tiernos provistos de hojitas tambidn se utilizan encurtidos como condimento. El uso de las alcaparras se remonta hasta la prehis- toria. Aunque la especie Capparis spinosa, que habita la regi6n mediterrdnea occidental es la mds utilizada, el subgFnero comprende 23 especies y subespecies que se extienden desde las costas Atldnticas hasta el Pacffico en Africa, Asia, Europa y Oceanfa. Se han registrado usos para 19 especies y subespecies, que son de interds alimentario y medicinal.

Key Words: medicinal plants; condiments; horticulture; food science; capers; Europe; West Asia; Central Asia; Oceania; Africa; indigenous knowledge; Capparis; Capparidaceae.

Capers and caper plants, wild or cultivated, are used in almost all of the range of Capparis subgenus Capparis, extending from West Africa to the Norfolk Islands in the Pacific. Most of the species are of alimentary or medicinal interest.

Current information of the uses of capers is extremely dispersed and refers to ambiguous sci- entific names; a review of the available data was undertaken in order to compare the past and pre- sent uses, and also to determine the possibilities for further development. We also present a re- view of both the local names and uses of the taxa of subgenus Capparis.

Capers of commerce are immature flower buds which have been pickled in vinegar or pre- served in granular salt. Semi-mature fruits and young shoots with small leaves may also be

J Received: 16 January 2002; accepted 31 January 2003.

pickled for use as a condiment (Alkire 2001). Existing national and international food legisla- tion define capers as the buds of just Capparis spinosa, but there is evidence that many others species are also used. The explanatory notes to the combined nomenclature of the European Community 98/C 287/01, published 15 Septem- ber 1998, define capers as the unopened flower buds of the spiny perennial shrub (Capparis spi- nosa) under class 0709 90 40. The FAO defines capers under the class number 0463--Other Vegetables--includes inter alia: bamboo shoots (Bambusa spp.); beets chards (Beta vulgaris); and capers (Capparis spinosa) (UN/ECE Statis- tical Division 2001). The European Commission (2001) includes capers in the list of Minor Crops, within Spices with Codex Classification HS 0773 and Bayer code CPPSP.

Capers were used in prehistoric times. Cap- paris spinosa was probably not utilized; how- ever, other species, such as C. aegyptia, C. or-

Economic Botany 57(4) pp. 515-534. 2003 �9 2003 by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

516 ECONOMIC BOTANY [VOL. 57

ientalis, C. zoharyi, and C. sicula, the common- est in the eastern Mediterranean and the Near East, may have been used. Caper seeds have been recovered from archaelogical sites in north- ern Syria: Mesolithic layers of Tell Abu Hureyra (Hillman 1975), Aceramic Mureybit (9th-8th millennium B.C.) (Zeist and Bakker-Heeres 1986), Pre-Pottery Neolithic A (8th-7th millen- nium B.c.) Jerf the Ahmar and Halula (Willcox 1996); Pre-Pottery Neolithic B (8th millennium B.C.) Djade (Willcox 1996). Also in Aceramic (8th-7th millennium B.C.) Aswad, site of Da- mascus basin (Zeist and Bakker-Heeres 1982). And in the Neolithic layers of Tell Abu Hureyra (Hillman 1975). And in the early 3rd millennium B.C., Tell Atij (McCorriston 1995). The Bronze Age jar containing carbonized flower buds and unripe fruits at Tell es Sweyhat, Syria, may in- dicate the established use of caper pickles in the Bronze Age (Zeist and Bakker-Heeres 1988). Campbell Thompson (cited by Blakelock and Townsend 1980) recorded that the dried leaves of caper steeped in vinegar have been used in Syria for application to ulcers and scabs of the head. Recently, 1999, we explored Syria and found C. aegyptia to be extremely abundant in the Damascus basin and C. sicula and C. aegyp- tia in the Upper Syrian Euphrates region. An exhaustive review of capers in Prehistory and Antiquity was published by Rivera et al. (2002).

MATERIALS AND METHODS

HERBARIA LABELS

The first source of information were the an- notated labels of herbarium specimens studied during our taxonomic revision of this subgenus. The herbaria consulted were BISH, BM, C, E, G, HUB, HUJ, JEPS, K, LINN, MA, MARSSJ, and MUB (acronyms from Holmgren, Holmgren and Barnett 1990). Data reported here as take from herbarium labels are indicated by the plant collector (and number, if any) in italics with the herbarium acronym for the herbarium in which we found the plant specimen.

FIELD NOTES

In our own field studies on the culture, uses, harvesting, and cultivars of Capparis in Moroc- co, peninsular Spain and the Balearic Islands, Syria, and Turkey, special care was taken to in- terview harvesters, farmers, and the staff of the local industries.

LITERATURE

We exhaustively surveyed the botanical, eth- nobotanical and agronomic literature. A search was also done on the world-wide web (WWW) in order to determine the relevance of capers in the current years.

COMPARISON OF INFORMATION

These data are compared for each taxon. Spe- cial care was taken to include local ethnobotan- ical studies. General, ambiguous references were excluded. The study area was defined by the dis- tribution range of subgenus Capparis, stretching from the Atlantic coasts of Africa and southern Europe, eastward to the Pacific Islands and Aus- tralia, southwestern Asia, Central Asia, Mada- gascar, and India.

BOTANICAL IDENTIFICATION AND NOMENCLATURE

Names and identifications of the plants from the other information sources have been adjusted this report to conform to those used by Inocen- cio (2001 ).

RESULTS Data are presented for individually for 19 taxa

and, finally, their cultivation, and commercial, food, and medicinal uses are summarized. We have not recorded names or uses for the follow- ing taxa and areas: Capparis mucronifolia Boiss. subsp, mucronifolia from Iran; Capparis mucronifolia Boiss. subsp, rosanoviana (B. Festsch.) Inocencio, D. Rivera, Ob6n, Alcaraz, from Tadjikistan; Capparis parviflora Boiss. subsp, sphaerocarpa Inocencio, D. Rivera, Ob6n and Alcaraz, from Afghanistan; and Cap- paris antanossarum Baillon, from Madagascar, either because data were not available to us or simply do not exist.

1. CAPPARIS SICULA VEILL. IN DUHAM. SUBSP. SICULA

This is the most widespread taxon, extending from the Atlantic coasts of southern Europe and North Africa to the eastern Mediterranean coast

Common Names: Spain: (Andalucfa) Alca- parrilla (flower buds), Alcaparrones (fruits), A1- caparra, Mata Panera, Tapanera (plant) (Mesa 1996; Martfnez, Gonzfilez-Tejero and Molero 1997); (Arag6n) Tapara (Femindez and Sanz 1993); (Castilla): Alcaparra; (Catalufia): Tfiper- es; (Murcia): Tapenera, Alcaparro, Tfipano, T~p-

2003] RIVERA ET AL.: FOOD AND MEDICINAL USES OF CAPPARIS 517

TABLE 1. RECOGNIZED CULTIVARS OF CAPPARIS SICULA, C. ZOHARYI, AND C. CORDIFOLIA.

Capparis sicula subsp, sicula

Cultivar Description Localities Reference

Spinoso, Flower buds less rounded, producing Salina, Italy Barbera and Di Lorenzo 1984; Spinoso flattened pyramidal capers, which Barbera et al. 1991; Fici and di Salina tend to flake. Leaves rounded, stout Gianguzzi 1997; Caccette

well developed spines. Often col- 1982 lected from wild populations.

It is becoming very rare due to the conspicuous presence of sharp thorns. Leaf tip also with a small thorn. Leaf oblong-ovate. Petiole long.

Decumbent twigs, stout well devel- oped spines. Pyramidal flower buds, good blossom but irregular. Large fruits.

Capparis zoharyt

Spinoso di Pantelleria, Italy Barbera et al. 1991; Fici and Pantelleria Gianguzzi 1997

Com~n, Del Murcia and Grana- Luna and P6rez 1985 Pais da, Spain

Cultivar Description Localities Reference

De Sail Flower buds rounded, producing glo- Sail, Morocco Inocencio 2001 bose capers, stout well developed spines.

Capparts cordifi)lia subssp, cordifolia

Cultivar Description Localities Reference

Alcaparras Flower buds rounded, producing glo- Guam, Marianas; Aklire 2001; Merrill 516, K bose capers, without well devel- Manila, Philip- oped spines, pines

ena, Caparra, (flower bud) Caparr6n, Mel6n (fruit) (Ob6n and Rivera 1991); (Valencia): Thp- era (fruit), Taperera (plant) (Ferr~ndez and Sanz 1993). France: Caprier. I taly: Capparo, Cap- pari; (Sicily): Cappero, Chiappara, Chiappara spinusa (Duhamel 1801; Williams 1971). Por- tugal: Alcaparras, Alcaparreira. Cyprus : Kap- pari (Anonymous 2001). Greece: Kappari; [the plant is here so common that many localities bear names derived from the abundant capers, viz. Kappari Beach in Myconos, or Kapparia in Korthi, or Kapparinos River]. Turkey: Kapari, Kebere, (Konya): Keditirnagi; (Gaziantep): Ke- here ~igegi (flowers), Kebera krkti (roots) (Bas- er, Honda and Miki 1986; 0zkahraman, Y~iksek and Kalkinma 2000).

Cultivation: Most of the uses are reported from the wild populations; the species is culti- vated as crop plant in Morocco, Italy, and Spain (Table 1).

Uses as Food: In the present, the use of this plant is centered in the flower buds, unripe fruits, and tender shoots, pickled in brine or vin-

egar and used as a condiment. European sources are Spain, Greece and Turkey. Cyprus : Shoots, unripe fruits, and flower buds are used in local salads "khoritiaki." They are preserved either in olive oil or in vinegar. Shoots with thorns, usu- ally pickled, are common in salads (Arnold 1985, Anonymous 2001). Spain: The flower buds, unripe fruits, and tender shoots are brined and preserved in vinegar in Murcia, Almerfa, and Granada. Murcian capers were exported as early as in 1478 (Zamora 1997). American com- panies are importing pickled caper and caper- berries from Lorca's local industries in Murcia, where they have been harvesting and processing capers and caperberries for decades (Wainer 2001). In Mfigina (Jarn) brined flowers and fruits are consumed as snacks or in salads (Mesa 1996). Greece : Capers have been used in Greece for thousands of years as a condiment. Today, capers are used in a wide variety of Greek dishes, from appetizers and poultry, to seafood (Nikolozakes 2001). Morocco: The flower buds are pickled. These are exported to

518 ECONOMIC BOTANY [VOL. 57

Fig. 1. Large containers with unprocessed brined C. sicula capers in Fes, Morocco. (Photograph by C. Inocencio).

the European Union and other countries by dif- ferent industries of the region of Fes and Mek- nes, Figs. 1 and 2 (Inocencio 2001; Lakrimi 2002).

Uses in Medicine: Turkey: One desert spoon- ful of powdered root with water is taken, after meals, for hemorrhoids (Baser, Honda and Miki 1986). Flowers are tonic, diuretic and blood pu- rifier in allergic diseases, an infusion of one spoonful in one liter water is taken one glassful after meals three times a day (Baser, Honda and Miki 1986). Fruits are externally applied for the treatment of headaches and piles in Anatolia (Oz- turk and 6zqelik 1991). Cyprus: The roots are boiled and used as an aperitif, tonic, astringent, antiemetic, anti-diarheal, anti-rheumatic. Leaves and fruits are used as aphrodisiac, anti-rheumatic, and anti-odontalgic (Arnold 1985). Italy: On the island of Lipari, a mixture powdered root barks of wild capers and walnut are used in the treat- ment of intermittent fevers (Estateolie 200l). In Sicily, leaves of the common caper, which have been gathered with the left hand, are ground up and applied to ulcers to make them suppurate (Williams 1971). Spain: The roots are used both as an anti-rheumatic and for tooth-ache in Gra- nada (Gonzgdez-Tejero 1989). Roots are used lo-

Fig. 2. Caper market in central Morocco. (Photograph by C. Inocencio).

2003] RIVERA ET AL.: FOOD AND MEDICINAL USES OF CAPPARIS 519

cally in Almerfa for baldness, tooth-ache, inflam- mations, and wounds in horses (Martfnez, Gon- zfilez-Tejero and Molero 1997). In Murcia, the root bark is used for tooth-ache and for healing wounds (Ob6n and Rivera 1991). Twigs are used in Almerfa for the treatment of ulcers (Martfnez, Gonzfilez-Tejero and Molero 1997).

2. CAPPARIS SICULA VEILL. IN DUHAM. SUBSP. HERBACEA (WILLD.)

INOCENCIO, D. RIVERA, OBON AND ALCARAZ (= C. herbacea Willd.,

C. ovata Desf. var herbacea (Willd.) Zohary)

Common Names: Armenia: (south) Aggeen- ko (fruit, Bobrov 1939); Kapar (Grosheim 1952). Georgia: (Georgian): Kapari (Grosheim 1952). Turkey: Keber (()zttirk and 0z~elik 1991). Russia: (Russian): Kapersi (Grosheim 1952). Azerbaidjau: (Azeri): Kever (Grosheim 1952), Turkmenistan: (Russian): Kapersi Ko- liutchie (Tcharieva 1988).

Uses as Food: Caucasus (Armenia, Azer- baidjan and Georgia): The flower buds of this plant are considered of nutrative. In the Cauca- sus they are consumed locally; in both the east- ern part of Ciscaucasia and in Daghestan, they are pickled by the small scale food industry. In southern Armenia, the ripe fruit is eaten raw (Bobrov 1939, 1970). Pickled fruit are used as condiment in Armenia and Azerbaidjan (Grosh- eim 1952). Uzbekistan: In the Fergana Valley the fruit is eaten raw; large quantities are alos stored for winter. Seeds are used for a source of oil (Bobrov 1939, 1970). Turkey: Flower buds are used as a seasoning in eastern Anatolia (Oz- ttirk and C)z~elik 1991).

Uses in Medicine: Turkey: Fruits are used externally for headaches and piles, in eastern Anatolia (Ozt~irk and Ozqelik 1991).

3. CAPPARIS SICULA VEILL. IN DUHAM. SUBSP. LEUCOPHYLLA (DC.) INOCENCIO,

D. RIVERA, OBON AND ALCARAZ (= C. leucophylla DC., C. spinosa L.

var. pubescens Zohary)

Common Names: Iran: Keverkai (J. Leonard, K); (Farsi): Mar Gir, Mar Gaz [= snake bite]; (Tehran): Rishah-i-kabar (Hooper 1937). Iraq: (Southern Desert) (Arabic) Shafallah; (Lower Jazira): Kabar; (Kurdistan): Kifri (Blakelock and Townsend 1980); (Persian) Kabar (bark) (Hoop- er 1937).

Uses as Food: Iran: Bakhtiari, the young fruit are pickled in vinegar. Throughout Iran the flower buds are harvested to make pickles (Hooper 1937). Iraq: The fruit are eaten by peo- ple, the plants are eaten by camels (Blakelock and Townsend 1980).

Uses in Medicine: Bahrain: Mahasneh, Ab- bas, and E1-Oqlah (1996) recorded the use in traditional medicine as a tonic and an expecto- rant. Iran: The plant is the source of many local herbal remedies for snake-bite, thus explaining the local Persian names Mar Gir and Mar Gaz (meaning snake bite) and others such as Alaf-i Mar (Blakelock and Townsend 1980). The light- colored root and the thick root bark are used in medicine, the bark being the Capparis Cortex Radicis of the old Persian Pharmacopoeia. They are both pungent and bitter and are given for intermittent fever and rheumatism (Hooper 1937).

4. CAPPARIS SICULA VEILL. IN DUHAM. SUBSP. MESOPOTAMICA

INOCENCIO, D. RIVERA, OBON AND ALCARAZ (= C. ovata Desf. var.

palaestina Zohary pro parte) Common Names: Saudi Arabia: Shafallah

(Mandaville 1990). Iraq: (Arabic) Kabar, Shaf- allah (Blakelock and Townsend 1980). Syria: (Bedouins): Shefellah (Rivera and Ob6n, MUB). Iran: Guh-i-kamar (= flower of the rocks), Alaf-i-Mar (= snake plant), Margaz (= snake bite); (Bakhtiari) Lagajee (Lee, K).

Uses as Food: Iran: the Bakhtiari tribe pro- cesses the unripe fruit in vinegar, for consump- tion (Lee, K). Syria and Turkey: Bedouins of the Syrian Upper Euphrates region harvest the flower buds for the Turkish food industry; it is not used as a food in Syria, but it is an item of commercial (Rivera and Ob6n, MUB).

Uses in Medicine: Iraq: The medicinal prop- erties of the root are still acknowledged by the Badawin: a decoction of the root taken orally as a palliative for rheumatism (Blakelock and Townsend 1980). Iran: It is used as a source of numerous medicinal remedies and for snake bites (Lee, K).

5. CAPPAmS SlCVt,4 VEILL. IN DUHAM. SUBSP. SINDIANA

INOCENCIO, RIVERA, OBON, ALCARAZ (= C. obovata Royle)

Common Names: India: Maratha language of Goa: Kabara (Dalgao 1898); Bombay: Kabar,

520 ECONOMIC BOTANY [VOL. 57

TABLE 2. RECOGNIZED CULTIVARS OF CAPPARIS ORIENTALIS AND C. XSPINOSA.

Capparis orientalis

Cultivar Description Localities Reference

Fulla Redona

Cavall

Nuciddara, Nocellara, ltaliana (in Spain)

Tondino

Testa di Lu- certola (=Lizard's head)

Ciavulara

Upper part of the shoots glabrous, un- Mallorca, Spain armed. Fruiting abundantly. Flower buds of superior quality.

Upper part of the shoots glabrous, un- Mallorca, Spain armed. Poor blossom.

Flower buds rounder, almost spheri- Pantelleria, Italy cal, considered of superior quality; extremely perfumed, orbicular leaves, notched, petiole short, ab- sence of thorns. It is the most widespread and productive. Product protected in Italy, lOP, since 27 December 1993.

Cultivated (a synonym of Nuciddara?) Salina and Pantel- leria, Italy

Flower buds less rounded, called "liz- Pantelleria, Italy ard's head" because of the charac- teristic lengthened pyramid shape of the capers. Petiole short, leaves ovate, no thorns. Considered of in- ferior quality. Its cultivation is con- tinually contracting.

Flower buds less rounded, flatter than Pantelleria, Italy those of Nocellara. Leaves ovate, more or less acute, petiole short, no thorns. Locally cultivated to a lesser extent. Product of inferior quality.

Capparis Xspinosa

lnocencio 2001, Luna and P6rez 1985

Luna and Pdrez 1985

Barbera and Di Lorenzo 1984; Barbera et al. 1991; Fici and Gianguzzi 1997

Caccette 1985

Barbera and Di Lorenzo 1984; Barbera et al. 1991

Fici and Gianguzzi 1997; Barbera et al. 1991

Cultivar Description Localities Reference

Josephine

Dolce di Filicudi e Alicudi

Nuccida, Nocella

Rosa

Variegata Figues Seques

One of the better Mediterranean se- lections but becoming rare.

Mild tasting flower buds.

Provence, France Alkire 2001

Eolie, Italy Alkire 2001

Flower buds rounded, considered of Salina, Italy superior quality. High productivity. Spines are usually present, particu- larly on the older branches, but not much developed. Leaves rounded, petiole long.

Prolific blossom. Blossom stems with- Mallorca, Spain out intercalary vegetative shoots. Not fruiting. Almost glabrous upper part of the shoots. Decaying leaves purple tinged. Flower buds of supe- rior quality.

Leaves striped yellow or white. France Poor blossom. Blossom stems with Mallorca, Spain

intercalary vegetative shoots. Flow- er buds large, flattened pyramidal, which tend to flake, thus of inferior quality. Sometimes bearing fruit. Upper part of the shoots almost glabrous.

Barbera and Di Lorenzo 1984; Fici and Gianguzzi 1997

Luna and Pdrez 1985; Rivera et al. 1999

Duhamel 1801 Luna and P~rez 1985; Rivera

et al. 1999

2003] RIVERA ET AL.: FOOD AND MEDICINAL USES OF CAPPARIS 521

TABLE 2. CONTINUED. XSPINOSA.

Capparis •

Cultivar Description Localities Reference

De las Prolific blossom. Blossom stems with- Mallorca, Spain Luna and Ptrez 1985; Rivera Muradas, out intercalary vegetative shoots, et al. 1999 Boscana Upper part of the shoots glabrous.

Redona

Peluda

Color6

Mallorquina, Fina

Fruiting abundantly. Flower buds of inferior quality.

Poor to good blossom. Flowering Mallorca, Spain stems with intercalary vegetative shoots. Flower buds rounded, of superior quality. Often bearing abundant fruit. Upper part of the shoots almost glabrous.

Prolific to normal blossom. Flowering Mallorca, Spain stems with intercalary vegetative shoots. Flower buds of superior quality. Often bearing fruit. Upper part of the shoots wooly.

Poor blossom. Flowering stems with- Mallorca, Spain out intercalary vegetative shoots. Flower buds of inferior quality. Not fruiting. Upper part of the shoots, almost glabrous.

Prolific blossom. Flowering stems Mallorca, Spain without intercalary vegetative shoots. Upper part of the shoots al- most glabrous. Decaying leaves yellow. Not fruiting. Flower buds of superior quality.

Luna and Ptrez 1985; Rivera et al. 1999

Rivera et al. 1999

Rivera et al. 1999

Rivera et al. 1999

Kalvhri (Cooke 1967); Kabar, Kawharg or Pa- netero (Jafri 1973). Pakis tan: (Baluchistan): Khwarg, Pahinro Kirap, Panetro Khafkhader (Baquar 1989).

Uses as Food: Pakis tan: camel and goat graze on the plant; fruit and buds are sometimes pickled for human consumption. The fruit is pickled in Punjab. Buds and unripe fruit are pickled as a condiment; ripe fruit is eaten raw (Baquar 1989; Edgeworth 1862). India: Young and ripe fruits are eaten raw; berries are boiled and eaten with curd. Unripe berries and flower buds are pickled. Leaves are used as fodder for goats, camels and sheep (Nasir and Rafiq 1995; Jafri 1973).

Uses in Medicine: India: An infusion of leaves is used in treating gonorrhoea. In Ladakh a decoction of leaves is drunk against hyper- acidity (Sundara 1993). Root bark is considered a diuretic, tonic, aperitif, expectorant, anthel- mintic, emmenagogue, analgesic; it is also used in paralysis, rheumatism, tubercular lymphade- nitis and enlarged spleen (splenomegaly).

Crushed leaves are used as a poultice for gout (Baquar 1989). Its used in Ayurvedic medicine as hepatic stimulant and protector, improving liver function (Alkire 2001).

6. CAPPARIS ORIENTALIS VEILL.

IN DUHAMEL (= C, rupestris Sm., C. spinosa L. subsp, rupestris (Sm.)

N y m a n )

Common Names: Spain: (Arag6n): Tapara. (Castilla): Alcaparra; (Catalufia): T~iperes. (Bal- earic Islands): T~tperes (flower buds), Gorrinets (fruits), Taparera (plant) (Guerau and Torres 1981). France: Caprier. I taly: Capparo, Cap- pari, Capperi (Galli 2001); (Sicily): Chiappara; (Pantelleria) Cappero (plants), Zucchette, Cap- perese, Cetriolini (island of Pantellaria, fruits) (Barbera and Di Lorenzo 1982); (Bologna): Ca- per, Caparen (Ungarelli 1985). Greece: Kappari. Portugal : Alcaparras, Alcaparreira. Libya: (Ar- abic) Cabbar, (Berber) Tilut (Jafri 1977).

Cultivation: On the island of Pantelleria, Italy (Table 2), although available in small quantities

522 ECONOMIC BOTANY [VOL. 57

from the wild, it is widely cultivated; the plants propagated from seed. Caper fields are placed close to the wild populations. Due to their prox- imity and insect cross pollination (nocturnal moths and mosquito), gene flow between the cultivated plants and the wild populations is high (Barbera and Di Lorenzo 1982; Fici and Gianguzzi 1997). The cultivar 'Nocellara' is predominant in Pantelleria, accounting for 90% of the local production. Flower buds are har- vested from May to October by the members of a local Cooperative Industry, involving 400 workers. Pantelleria's 'Nocellara' cultivar re- ceived the status of Indicazione Geographica Protetta from the Italian authorities (Verde-Oggi 2001; Galli 2001). The species is grown as or- namental in the Balearic Islands of Spain, France, and Italy, where it is sold as Capparis spinosa 'Inermis'.

Uses as Food: Greece: In the Aegean, fruit are pickled; in Candia, leaves and flower buds are pickled. Spain: Tender fruit (gorrinets) and flower buds (t~peres), brined and pickled, are used in traditional foods of Ibiza in the Balearic Islands (Guerau and Torres 1981). Italy: Fruits are known in Italy as zucchette, capperese and locally in Pantelleria as cetriolini. The fruits are not gathered every year; for this purpose part of the flower buds are allowed to reach anthesis and to produce fruit. Harvest of flower buds and fruits are mutually exclusive, and normally flow- er buds are much prized (Barbera and Di Lor- enzo 1982). Pliny the Elder, in his Natural His- tory, written in the 1st century A.D. described the "seed" as a well-known article of food. The pickled fruit are said to have a peppery flavor. Capers are used in numerous recipes of the Ital- ian cuisine: capers are an essential ingredient in Pizza Napoletana and Pizza alla Siciliana. They are also used in Salsa Verde for Spaghetti alla Puttanesca, Galantina di Pollo, and Penne alla Sorrentina (Galli 2001). Libya: Young flower buds and fruits are pickled (Jafri 1977).

Uses in Medicine: Libya: The tea made with the plant is used in Libya as a remedy for stom- ach ailments (E1-Gadi and Bshana 1986; Trotter 1915). It has been investigated for antitumor ac- tivity (Duke 2001).

7. CAPPARIS XSPINOSA L.

Common Names: Spain: (Arag6n): Tapara. (Castilla): Alcaparra. (Catalufia): T~iperes. (Mur- cia): Tapenera, Tapena, Alcaparro, Caparro, T~i-

pano. Italy: Capparo, Cappari; (Sicilia): Chiap- para. Portugal: Alcaparras, Alcaparreira. France: C~prier (plant), C~pres (flower buds), Cornichons (fruits) (Cazin 1868); (Provence): Tapari6, Tapenir, Taperi6. (Toulouse): Capriai (Reguis 1878; Rolland 1967).

Cultivation: It is rarely found in the wild, which are populations escaped from cultivation. The wild plants commonly referred as C. spi- nosa belong either to C. sicula or C. orientalis. Italy: In the 1990s the Italian production was concentrated on two volcanic islets in the vicin- ity of Sicily: Pantelleria in the Strait of Sicily and Salina in the Tyrrhenian Sea, a total surface of ca. 950 ha (Fici and Gianguzzi 1997). Root cuttings are used on propagation on the Aeolian archipelago (Italy) and in Mallorca (Spain). When hybrids of this taxon are propagated by seed, the resultant progeny are highly diverse. In this case, the segregate types can be propagated clonally by cuttings. Hybrid plants can be rec- ognized by their parental combinations of leaf shapes, and very often by the presence of dif- ferent foliage colors, reddish, deep green, yel- lowish, etc. (Barbera and Di Lorenzo 1984; Ri- vera et al. 1999; Inocencio 2001). Spain: In Campos, Felanitx, and Llub/, it has been culti- vated extensively. During the last century (1860s to 1960s), harvest of flower buds was an almost exclusively a woman's task (Segura 1997).

Uses as Food: Flower buds, fruits, and, lo- cally, tender young shoots (including immature small leaves), pickled in brine or vinegar as a condiment. European sources are the Balearic Islands of Spain, Italy, and Provence in France (Table 2).

Uses in Medicine: Capers are said to reduce flatulence and to be anti-rheumatic (Alkire 2001). France: Flower buds and root bark are used for cleansing the liver and the spleen, roots are used as a tonic and a diuretic (Cazin 1868).

8. CAPPARIS OVATA DESF. (= C. fon tanes i i DC.)

This species is an African endemic which is often erroneously cited from Italy and Spain (Barbera and Di Lorenzo, 1984), because of confusion with C. sicula Veill.

Common Names: Algeria: (Berber Tema- check) Talulut, Telulut, Telofilofit, Touloulout (Benchelah et al. 2000; Chevalier 1938); (Ara- bic) Khabbar, Soukoum (Keith s.n., K; H. Du-

2003] RIVERA ET AL.: FOOD AND MEDICINAL USES OF CAPPARIS 523

veyrier P 130676); Felfel-el-gebel (Biskra, Al- geria) (Schweinfurth 1912). Chad: (Toubou): Gozui, Gozou, Kozohou (H. Broo, 52 K; Che- valier 1938). Morocco: (Arabic) E1 Kabbar, (Berber) Teililout, Teiloulout (N~gre 1961).

Uses as Food: Chad: The women of the Ti- besti mountains harvest the fruit and mash them with milk to make a gruel to consume as a food (St. Serole 57, P 130669; Chevalier 1938); Fod- der for gazella. Algeria: The Tuaregs of Tassili region use only the flower buds as capers, the Tuaregs of the Hoggar do not use the species (Benchelah et al. 2000; Gast 2000).

Uses in Medicine: Algeria: According to Maire (1933) it is used in the Hoggar as anti- inflammatory, for headache, and for stomach ache. Crushed leaves are used in the treatment of "la gale des chameaux," and after boiling they are applied for lumbago (Benchelah et al. 2000; Gast 2000). Morocco: In Tissint the pow- dered dry root is used for blenorhagy. A mixture of caper fruits with leaves of Waronia saharae Benth. and Coss. and olive oil is used for the common cold. Boiled fruits alone or mixed with Waronia are used for stomach ailments. In the Sahara and Dra regions fruits and leaves, mixed with Cleome arabica L. subsp, amblyocarpa and olive oil, are used as anti-inflammatory oint- ments (Bellakhdar 1997)

9. CAPPARIS HEREROENSIS SCHINZ.

Common Names: unknown. Uses as Fodder: It is eaten by mules and cam-

els but horses do not do so (Watt and Breyer- Brandwijk 1962).

Uses in Medicine: The root seems to be well- known as poisonous in the Wankie district, Zim- babwe, Africa; the people sprinkle the powdered root on meat to kill lions and leopards (Watt and Breyer-Brandwijk 1962). Both of the uses men- tioned above are doubtful because they are re- ported from the Wankie district of Zimbabwe; this species is endemic to Namibia

10. CAPPARIS AEGYPTIA LAM. (= C. spinosa L. vat. deserti Zohary)

Common Names: Egypt: (Arabic) Kabar, Quabar, Lassaf, Latsst.ssaf, Shafellah (Rolland 1967; Schweinfurth 1912); (Coptic): Kemegeoc, Kemapic. Iraq: (Diwaniya, Ba'quba, Southern Marshes): Kabar; (Basra): Shafallah (Blakelock and Townsend 1980).

Uses as Food: Egypt: The seeds are added to

wine to keep it sweet and are also used as a condiment (Renfrew 1987). Iraq: Pickles are made from fruits in Basra (Blakelock and Town- send 1980).

Uses in Medicine: Egypt: Powdered root bark is mentioned by Prospero Alpino (16th century A.D.) as a treatment for skin diseases, as a verm- ifugue, an antitummoral, and an emmenagogue (Fenoyl 1980). In Egypt, it is used medicinally by the Copts to treat wounds (Manniche 1989).

11. CAPPARIS ZOHARYI INOCENCIO, D. RIVERA, OBON AND

ALCARAZ Common Names: Spain: (Murcia) Alcaparra.

Morocco: Kabar. Syria: (Bedouin): Shefellah (Rivera and Ob6n, MUB); (Aleppo) Hasiset el- qebbar, Zaheg qabbar (plant), Qebbar (flower buds), Qabbar (roots) (Honda, Miki and Saito 1990).

Cultivation: Most of the uses are reported from material harvested from the wild popula- tions, but the species is locally grown as crop plant around Sail, Morocco (Table 1).

Uses as Food: Morocco: In the region of Sail, the flower buds are harvested for pickling and consumption (Inocencio 2001). They are ex- ported to the European Union and other coun- tries by industries of the Casablanca and Mar- rakech region (Lakrimi 2002). Syria: A jar con- taining flower buds and unripe fruits at Tell es Sweyhat may indicate the use of caper pickles in the Bronze Age (Zeist and Bakker-Heeres 1988).

Uses in Medicine: Mauri tania : Cooked leaves in a pomade are used on camels for treat- ment of skin diseases and external parasites (Lehmann 2001). Syria: Campbell Thompson (cited by Blakelock and Townsend 1980) re- corded that dried leaves steeped in vinegar are used in Syria for application to ulcers and scabs of the head. Flower buds and roots are sold in Aleppo for medicinal purposes (Honda, Miki and Saito 1990). Rauwolff noted in Aleppo that "such plenty of capers that they are not at all esteemed" growing on old walls, the natives pickling their flowers before they open (Camp- bell-Thompson 1949).

12. CAPPARIS ATLANTICA INOCENC10, D. RIVERA, OBON AND

ALCARAZ Common Names: Morocco: Kabar (Inocencio

2001).

524 ECONOMIC BOTANY [VOL. 57

Uses as Food: Morocco: As a condiment, flower buds and fruits are prepared pickled in vinegar or brine. In the mountains of the High Atlas, the flower buds are harvested for pickling and consumption (Inocencio 2001). These are exported to the European Union and other coun- tries by industries of Marrakech region (Lakrimi 2002).

13. CAPPARIS PARVIFLORA BOISS. SUBSP. PARVIFLORA

(= C. leucophylla DC. vat. parviflora (Boiss.) Zohary)

Common Names: Iran: (Farsi): Mara Gira, Mar Gir (Blakelock and Townsend 1980). Iraq: (Kurdish): Halabja; (City of Kifri): Kabaruk, (Blakelock and Townsend 1980).

Uses as Food: Iran: The fruits are eaten (Blakelock and Townsend 1980).

Uses in Medicine: Iran: Persian healers use boiled roots in olive or sesame oil and white wine for spleen diseases associated with inter- mittent fever (Blakelock and Townsend 1980).

14. CAPPARIS PARVIFLORA BOISS. SUBSP. KUROICA (ZOHARY)

INOCENCIO, D. RIVERA, OBON AND ALCARAZ.

Common Names: l raq: (Kurdish) Halabja, (Blakelock and Townsend 1980).

Uses as Food: Iraq: The fruits are eaten (Blakelock and Townsend 1980).

15. CAPPARIS INERMIS FORSSKAL. (= C. cartilaginea Decne., C. galeata

Fresen.)

Common Names: Egypt: (in Sinai): Arabic names Felfel-jibbel, Goah, Goah-kulul, Latss.tssaf, Lassaf (Schweinfurth 1912). Somalia: Goah, Goah-Kulul, Goh, Gombor (Thulin 1993). Kenia: Chepkogh, Chepteretwa, Gorra, Ilngorochi, Lea- char, Lokapilak, Mtunguru, Mbaruti, Olatunde, Qadhu (Beentje 1994; Dale and Greenway 1961; Lehmann 2001). Iran: (Balouchistan) Gorilimbuk Blatter (Parsa 1951). Saudi Arabia: Lassaf (BoyS, K), Lusef (plant), 'Aslub, 'Albelib (fruit) (Ghan- zafar 1994); Lfitssaf (South Arabia) (Schweinfurth 1912); Nutss~if (Hogela, South Arabia) (Schwein- furth 1912). India: Karat (Saurashtra K). Dhofar: (Jibbali names): L6sef, Aselib (fruit). (Dhofari Ar- abic): Lfisfeh (plant), Albelib, Aslu~b (fruit) (Miller and Morris 1988). Yemen: Lattssaf (Forss-

kill cited by Schweinfurth 1912); La~af, Ni~af (Wood 1997).

Uses as Food: Egypt: The people of the Sinai uses capers as a condiment, known as mountain pepper. The fruit are eaten as food by the Bis- charin and Kushmaan Bedouins. Dried fruits are the main ingredient of a nutritious, spicy drink called mariida, which can be kept in a waterskin for 1 to 2 months (Goodman and Hobbs 1988). Dhofar: The fruit is consumed for its sweet pulp. The fruit, once ripened, turns a beautiful scarlet; the flesh inside is sweet and delicious. The bitter skin is discarded. The fruit is high in vitamin C and contains significant amounts of protein and carbohydrate. The foliage was tra- ditionally an important fodder for livestock. Sickly camels and goats were herded to areas where these plants proliferated and were en- couraged to browse the bushes. The severe bouts of diarrhea that initially followed caused the an- imals to deteriorate in condition, but after a time, their appetite returned and they grew fat and sleek and their milk increased in quantity and quality (Miller and Morris 1988). Somalia: Fruits are eaten raw (Thulin 1993).

Uses in Medicine: Egypt: A poultice to com- bat rheumatism is made by cooking fruits with salt and vegetable oil, letting the mash cool, and then applying it to the painful area (Goodman and Hobbs 1988). Oman: Water from boiled leaves was used for washings after child-birth, as a remedy for pains and as an antiseptic. Wa- ter, from boiling shoots and leaves, is employed in the treatment of skin inflammation, bruises and also as a remedy for pain and inflammation in the case of snake bites. A paste is prepared with the leaves and shoots and externally ap- plied for inflammation in joints, sprains, mus- cular contractions, tendinitis, paralysis of the body members, rheumatism, and problems of the knees. An identical paste is used to the cure of livestock infected with parasites and ticks. It is used against ticks by making them come off by applying to the wound the leaves or their juice, when the tick is still in the skin (Miller and Mor- ris 1988). Saudi Arabia: A decoction of leaves and stems is used for bruises, childbirth, ear- ache, headache, paralysis, snakebites, and swell- ings (Ghanzafar 1994). Kenia: A decoction of leaves is used either in ocular bath or as eye lotion for eye diseases for humans and cattle. A decoction of leaves is used for indigestion. Leaves, twigs, and stems are chewed for colics.

2003] RIVERA ET AL.: FOOD AND MEDICINAL USES OF CAPPARIS 525

Fig. 3. Different grades and types of capers. (Photograph by C. Inocencio).

Juice of the root is used for dermatitis and skin ulcers (Lehmann 2001). Tanzania: Crushed stems and leaves are given in an infusion for cattle fevers by the Masai (Lehmann 2001). So- malia: The leaves are chewed to treat coughs (Thulin 1993). In this same area, dried leaves mixed with wood ashes are used as a sort of chewing compound; leaves are also used as a medicine to cure the common cold in new born goats. Around twenty leaves are blended with water and then are crushed in a cup. This quan- tity is enough to treat two young goats. (Peck 226, K).

16. CAPPARI$ NAVAULENStS DC. (= C. himalayensis Jafri).

Common Names: India: Karil (Collet 1902); Kabra (Sundara 1993). Pakis tan : (Punjab): Kakri, Kander, Kabra (Baquar 1989); (Sanskrit): Kakadani; (Hindi or Hindustani and Punjabi) Kabra (Nadkarni 1976).

Uses as Food: fruits and flower buds are used as a condiment.

Uses in Medicine: India: According to Kak- rani and Saluja (1994) in Kutch district, Gujarat State, leaves are used as an antirheumatic. The root bark is bitter, hot and dry; aperient, tonic,

expectorant, antihelminthic, emmenagogue, an- algesic; good in treating rheumatism, paralysis, toothache, tubercular lymphadenitis, and en- larged spleen (splenomegaly); the juice kills worms in the ear (Yunani) (Kiritikar and Basu 1987). In Ormara and Las Bela, a jelly is pre- pared from fruits to cure rheumatism and snake bites (Hughes- Buller cited by Kiritikar and Basu 1987). Much juice is obtained from crushed berries, but, if they are dry, a small amount of water can be added, and the cold juice is poured into the ear as a cure for ear-ache (Hot- son cited by Kiritikar and Basu 1987). Pakistan: Used in palsy, dropsy, gout and rheumatism (Nadkarni 1976). Dried bark of the root is con- sidered a diuretic. In Kangra (Punjab), macer- ated roots are applied to sores (Kiritikar and Basu 1987).

17. CAPPARIS CORDIFOLIA LAM. SUBSP. COROWOLtA ( = C. mariana DC.)

Common Names: Solomon Islands: (Island of Santa Ana): Larafaku (R. Maurias 17218, K). Archipelago of Tuamotu (Island of Malekula): Puapila (J. Florence 3034, K). (Niau): Tuipiro- piro. Cook Islands (Mitiaro): Papiro (C. Luttrel 8, K). Fijl Islands (Islands of Rotuma and Mea-

526 ECONOMIC BOTANY [VOL. 57

TABLE 3. COMMERCIAL GRADES OF CAPER FLOWER BUDS. DATA FROM BARBERA AND DI LORENZO

(1984); BACCARO (1978); CAPC (2001); LUNA AND P~REZ (1985). A PRIMmVE CLASSIFICATION IN "NOMPAREILLE, CAPUCINE, CAPOTE, SENCODE AND TROISIEME" IS GIVEN BY DUHAMEL (1801) .

Diameters Spain . Spain Italy Italy France Italy (caliper) commercial industry industry commercial (Pantelleria)

C. • and C. sicula 0-7 mm Nonpareil les Primeras Grado 1 C a p p e r i n i Nompareilles Piccoli 7-8 mm Surfines Segundas Grado 2 Capperini Surfines Medi 8-9 mm Capucines Terceras Grado 3 Capperini Capucines Medi 9-10 mm Capotes Cuartas Grado 4 Capperi Capotes Grosso

10-11 mm Capotes Cuartas Grado 5 Capperi Fines Grosso 11-12 mm Fines Quintas Grado 6 Capperi Grosses Grosso 12-13 mm Fines Quintas Grado 7 Capperi Grosses Grosso 13-14 mm Gruesas Gruesas Grado 8 Capperi Grosses ? 14-15 mm Gruesas Gruesas Grado 9 Capperi Grosses - - 15-16 mm Gruesas Gruesas Grado 10 Capperi Grosses - - 1 &more Gruesas Gruesas Grado 11 Capperoni Grosses - -

mea): Periro, Piriro (H. St. John 19366, BISH 615083). Hawaii: Maiapilo or Puapilo. Niue Is- land: Pamoko. Archipelago of the Marianas (Island Cocos): Acparas, Acaparas, Atkaparas, name of Spanish origin, (B.C. Stone s.n., BISH 135459); (Island of Guam): Alcaparra (W.E. SaC-

ford and A. Seale 1108, US 516110). Islands Salomrn (Venmauis): Nefuerrfuerr; (Vorfar, Tisbel): Bangforrlam, (R. Maurias,17218, K). Island of Nauru: Ekobobwija (A. Burge, 17, K). Rurutu (Tubuai): Tiare-teina-mato. (Zepernick 1972)

Cultivation: It is generally used from the wild but there are two references of material being used from locally cultivated plants (Table 1). Marianas: Formerly grown as a commercial crop in Guam (Alkire 2001). Philippines: Ac- cording to a specimen (Merrill 516, K), this spe- cies was introduced into the Philippines from the island of Guam, being grown in the towns of Forfiaque and Malab6n, near Manila, and per- haps elsewhere. According to Jacobs (1965), it was introduced in the Pacific islands either by the Spaniards or the Portuguese, from southern Europe (citing erroneously Merrill 516, K). On the island of Guam, an ancient colony of Spain, the common names of Spanish origin are still used. This species is not present in Europe nor are any of the remaining taxa belonging to Sec- tion Galeatae. On morphological grounds the nearest species is C. antanossarum Baillon, from Madagascar.

Uses as Food." Marianas and Philippines: Pickled flower buds are used as condiment.

Uses in Medicine: Vanuatu: It has some un- specified medicinal uses in Vanuatu (P. Curry 1403, K).

18. CAPPARIS CORDIFOLIA LAM. SUBSP. SANDWICHIANA ( D E . )

INOCENCIO, D. RIVERA, OBON, AND ALCARAZ (= C. sandwich iana DC.)

Common Names: Island of Niue: Pamoko (T.G. Yunker 10112, K). Archipel of Hawaii: (island of Hawaii) Puapilo (G.O. Fagerlund, A.L. Mitchell 885, K; Nagata 1971; Wagner, Herbst and Sohmer 1990; Zepernick 1972), Maiapilo (H. St. John 22463, US 2446980; Wag- ner, Herbst and Sohmer 1990; Zepernick 1972); (island of Maui): Pilo (K. Livermore s.n., BISH 405983; Wagner, Herbst, and Sohmer 1990; Ze- pernick 1972). Archipelago of Tuamotu: (Is- land of Niau): Tupiropiro (J. Florence 10044, P ] 30657), Tufirofiro (Zepernick 1972).

Uses as Food." It is presumably consumed but it was neither explicitly stated on the herbarium labels nor in the literature.

Uses in Medicine: Hawaii: It is used to repair broken bones, aching backs and muscles by the akamai, the Hawaiian priest of herbal medicine (Bornhorst 2000; Duke 2001; Nagata 1971; Ze- pernick 1972).

19. CAPPARISNUMMULARIA DC.

Common Names: Australia: (English) Wild Passion Fruit, Flinders Rose, Split Arse, Split Jack, and Nipan (Ronald de Fossard pers. comm.); (tribal names): (Alyawarr) Arrwerneng,

2003] RIVERA ET AL.: FOOD AND MEDICINAL USES OF CAPPARIS 527

Fig. 4. Industrial screening of capers in Fes, Mo- rocco. (Photograph by C. lnocencio).

(Anmatyerr) Arratning, Arrweneng; (Eastern Arrernte) Arrutnenge; (Western Arrernte) Ratn- inge; (Pintupi) Wraningi; (Warlpiri) Minkilyan- anga (Latz 1995); (Western Australia Territo- ries): (the flowers) Bajalor or Bajala (W.A. Mich- ell s.n., K).

Cultivation: All uses refer to wild popula- tions. Australia: Jacobs (1965) cites that C. nummularia is lacking in the earliest collections of Australian plants; thus its introduction to Aus- tralia seems to have occurred before the defini- tive establishment of the British colony, some- where in the Northwestern coast, extending later towards central Australia and Queensland. The only western taxon presumably ancestor to this species is C. antanossarum Baillon from Mad- agascar, Aldabra, and Island Europe. But G. Chippendale and Greenway (cited by Jacobs 1965) note that C. nummularia is a true native in the central semi-arid areas of Australia, form- ing small patches, in the gorges of the mountains and on river beds.

Uses as Food: Australia: Edible (A. C. Boyle s.n., K). Ripe fruits of this shrub are harvested by Australian aborigenes, who consume the pulp (Ronald de Fossard pers. comm.; Latz 1995).

Uses as Fodder: Fodder (Wewson 1982).

DISCUSSION

USES FROM THE WILD

Most of the traditional and commercial uses of Capparis depend exclusively on the collec- tions from the wild and thus fall within the cat- egory of Non-Wood Forest Products (NWFP) as defined by the FAO. For instance, wild Capparis accounts for 17% in the export of NWFP in Tur- key (Koc, Aksu and Kurtoglu 2000).

CULTIVATION

Cultivation has been only reported for C. Xspinosa, C. orientalis, C. sicula, C. zoharyi and C. cordifolia (Tables 1 and 2). Capparis xspinosa is a hybrid C. sicula and C. orientalis. The hybridization of these two species is fre- quent in the wild; there are many intermediate types in cultivation. This hybridization produces variable, sexually reproducing populations of cross-pollinated plants. Mass selection is man's oldest plant breeding method, it is also one of the best plant breeding methods to enhance germplasm and develop improved cultivars to- day. Most of the cultivars are within C. Xspi- nosa (Barbera and Di Lorenzo 1984; Rivera et al. 1999). Due to proximity and entomophilous pollination, gene flow between cultivated plants and wild populations is high within the complex C. Xspinosa, C. sicula, and C. orientalis (Bar- bera, Di Lorenzo and Barone 1991). Propagation of the species mentioned above (specially C. Xspinosa) can be result from both seed and root- ed cuttings. In the first case, seeds gathered from the dehiscent fruits when they have ripened to- wards the end of summer are sown at a depth of a few centimeters in open field seedbeds in the month of February. About 5% of these seeds germinate in April or May, the other 95% are unpredictable in germination, although some 5% more germinate within the next year. The plants obtained from seed are heterogeneous in terms of leaf shape, thorns shape, and flower mor- phology; however, they are considered homo- geneous enough for the purposes of quality and quantity requirements of the crop. Therefore, it is common to grow several strains together that may be recognized as distinct by the farmers. The seed propagation method is commonly used on Pantelleria Island, Italy, for C. orientalis 'Nucciddara'.

The use of rooted cuttings is common in the

528 ECONOMIC BOTANY [VOL. 57

Fig. 5. Commercial samples of pickled capers from Italy, Spain, Turkey, Morocco, and Greece. (Photograph by C. Inocencio).

Aeolian archielago, Italy, and in Mallorca, Spain, for C. Xspinosa cultivars. To obtain a sat- isfactory percentage of rooting, which is never higher that 50%, it is necessary to take cuttings at the moment of pruning, in February, March, or April, from the most vigorous branches of a diameter not less than 1.5 cm (Barbera and Di Lorenzo 1984). Watering of the cuttings by sub- mersion for one month is also done in Italy pre- vious to planting (Baccaro 1978). In the XVIth century De Serres (facsimile edition of 1996) recognized cuttings as the best method for prop- agation of capers. Capers were widely grown in Provence (France), especially in the region of Toulon. Propagation by rooted cuttings or seeds in autumn and protected in the winter with mulch, was the usual practice (Cazin 1868).

In Italy, plants are spaced 2 to 2.5 meters apart. Caper fields may have about 1500 to 2500 (3000) plants per hectare. Under Italian legisla- tion, only 1500 are allowed for C. orientalis 'Nuciddara' fields on Pantelleria island. Seed-

grown plants or rooted cuttings may start pro- ducing flowers after their first to fourth years, depending on the soil conditions and water availability. The full yield is obtained after the fourth year. Each year pruning takes place in winter, to eliminate the dead wood and to reduce the live branches to a length of 1 cm. Only the first year branches flower. The cultivation, till- age, and fertilizing operations are carried out regularly (Alkire 2001; Barbera and Di Lorenzo 1984; Cacette 1985; Lettieri 2001).

The average yield of flower bud per year is ca. 1-1.5 kg for C. orientalis 'Nuciddara' on Pantelleria and ca. 2-3 kg for C. Xspinosa and C. sicula cultivars in Salina, Italy. The yield per hectare reaches between 3 to 4 tons for for C. Xspinosa and C. sicula cultivars and 2.2 tons for C. orientalis 'Nuciddara' (Lettieri 2001). Production spreads over a long period, begin- ning in May and lasting to the end of August. There is a tendency to leave very short gaps of time in-between each picking, as the best quality

2003] RIVERA ET AL.: FOOD AND MEDICINAL USES OF CAPPAR1S 529

Fig. 6. Pickled unripe fruits of C. sicula from Spain (Photograph by D. Rivera).

flower buds are the smallest. There are about 9 to 12 harvests each season (Barbera and Di Lor- enzo 1984).

COMMERCIAL USES

For commercial purposes, the grades of caper are based on the bud diameter, ranging from 7 to 27 mm. The commercial names are nonpar- eilles, surfines, etc. (Table 3, Fig. 3). The smaller grades are preferred. On Pantelleria, the largest grade is for C. orientalis 'Nuciddara' is only 14 mm, with 60% below 12 mm, which justifies that 'Nuciddara' capers are considered of best quality (Barbera and Di Lorenzo 1982, 1984). The average diameter is 9 mm (Lettieri 2001). The differences in bud diameters mean results in harvest difficulties. There are ca. 7000 flower buds/kg of nonpareilles (Table 3), 4000 buds/kg of surfines or capucines, ca. 2000 buds/kg of cappottes, ca. 1300 buds/kg of fines, and only ca. 800 buds/kg of grosses (Luna and Prrez 1985). It takes ten times more to harvest one kilogram of the 7 mm grade than one kilogram

of the 14 mm grade. The flower buds are screened through mesh and then classified by sizes (Fig. 4). The different grades are processed in brine (Fig. 1), and one month later vinegar is added, or, alternatively, they are preserved in salt. The salt procedure is preferred by gastron- omists, as it is better at preserving the strong flavor of the capers. The flower buds are covered and mixed with sea salt at 40% of their crude weight, inside wooden barrels and are kept for ten days. The brine from the mixture of water from the capers themselves and the sea salt helps the maturation of capers (Lettieri 2001).

USES AS FOOD

Akgtil and C)zcan (1999) found that the seeds of capers (C. sicula and C. Xspinosa) are rich in protein, oil, and fibre. The high proportion of unsaturated fatty acids suggests that they might be valuable as food, and therefore, may have been used as such in the past. Even though the use as food of capers is restricted to pickled flower buds (Fig. 5), unripe fruits (Fig. 6), and

530 ECONOMIC BOTANY [VOL. 57

Fig. 7. Pickled young shoots of C. sicula from Spain (Photograph by D. Rivera).

shoots (Fig. 7) in modern times, it seems likely that consumption of the ripe fruit, also with sweet pulp, played a relevant role in ancient us- age and in the beginning of the cultivation and domestication of capers. This use is recorded for fruits of C. sicula subsp, herbacea in Armenia and Uzbekistan, C. sicula subsp, leucophylla in Iraq, C. sicula subsp, sindiana in Pakistan and India, C. ovata Desf. in Chad, C. parviflora subsp, parviflora in Iran, C. parviflora subsp. kurdica in Iraq, C. inermis in Egypt, Dhofar and Somalia and C. nummularia DC. in Australia.

MEDICINAL USES

Medicinal uses of capers have been reported at least since the 1st century A.D., in texts of Dioscorides, Pliny, and others. Those are sum- marized in Table 4. The medicinal use, espe- cially of caper roots and bark, persisted during the Middle Ages, as confirmed by Celsius, Al- exander of Tralles, The Antidotary of Cam- bridge, The Antidotary of Reichenau, Marcellus,

Oribassus, Celsus, Scribonius, Paul of Egina, and others (Opsomer, 1989).

Capers are extremely rich in phenolic com- pounds as quercetin 3-rutinoside, kaempferol 3- rutinoside, and kaempferol 3-rhamnosyl-rutino- side; free aglycones as quercetin and kaempferol are produced during the brining process. Brining extracts most of the kaempferol 3-rhamnosyl ru- tionside present in the original flower buds. A 10 g serving of capers will provide 30-160 mg of flavonoid glycosides, at least the same amount of quercetin as 100 g of onions yield (Inocencio et al. 2000). Capers contain thioglu- cosides (known as glucosinolates) which release isothiocyanates (mustard oils) when the plant is crushed. Typically the plant yields methyl iso- thiocyanate from methyl glucosinolate, also known as glucocapparin. These mustard oils are responsible for the pungent flavor typical of ca- pers (Capparaceae) and the mustard family (Cru- ciferae), but they also have skin irritant effects and may also cause contact allergies (Mitchell

2003] RIVERA ET AL.: FOOD AND MEDICINAL USES OF CAPPARIS 531

TABLE 4. MEDICINAL USES OF CAPERS IN GRECO-ROMAN CULTURES. DATA ACCORDING TO DIOSCORIDES AND PLINY (1ST. CENT. A.D.), AND PALLADIO (IVTH CENT. A.D.) (GUNTHER 1968; DESFONTAINES 1829A, 1829B; MOURE 1990).

Uses Part of plant Authors

For the spleen Good for the spleen, as emetic and

diuretic Expels urine Expels the bloody excrement Expels the tape worms Help the sciatica Good for ruptures and convulsions Drive out the menstrual discharge Drives away the mucus from the head For mouth ulcers Cleanse ulcers Analgesic for pained teeth Takes away white vitiligo spots Dissolve hardnesses Kill the worms in the ears or as

analgesic for ear ache To keep healthy the oxen Those that daily eat capers are protect-

ed from palsey and spleen ache It is not useful for stomach ache

Fruits, root bark Dioscorides, Pliny Root bark in wine Pliny

Fruits, root bark Fruits, root bark The plant in vinegar and honey Fruits, root bark Fruits, root bark Fruits, root bark Fruits, root bark Plant boiled in vinegar Root bark and leaves with honey Fruits, root bark chewed or seeds in vinegar Frictions with bruised root bark Roots and leaves Roots and leaves boiled in oil

Dioscorides Dioscorides Pliny Dioscorides, Pliny Dioscorides Dioscorides Dioscorides Pliny Dioscorides, Pliny Dioscorides, Pliny Dioscorides, Pliny Dioscorides Dioscorides, Pliny

Bruised caper leaves, myrtle, and cypress Palladio Flower buds Pliny

The plant Pliny

1974). The following species have been reported to contain glucosinolates (Ahmed et al. 1972): C. aegyptia, C. inermis, C. sicula subsp, meso- potamica, C. orientalis.

Khakberdyev et al. (1968) described experi- ments involving oral administration of a prepa- ration of the root of C. sicula subsp, herbacea to guinea pigs, prior to innoculation with a sub- cutaneous injection of various animal and plant allergens to which the guinea pigs had been sen- sitized. In all cases, the animals treated with the Capparis preparation suffered no ill effects but untreated guinea pigs died of anaphylactic shock (Schmidt 2001).

CONCLUSIONS

In this paper we have presented data concern- ing the uses and common names for the species of Capparis subgenus Capparis the uses being fundamentally alimentary or medicinal. Our in- terpretation of the data leads to the following conclusions.

1. The general use of flower buds, fruits and shoots, preserved in different ways (salt, brine, vinegar), is as food. Capparis orien- talis and C. • are most commonly

used in commercial samples, but also C. zo- haryi and C. sicula are relatively frequently encountered.

2. Ripe fruits have been consumed for their seeds and sweet pulp.

3. Medicinal uses involve mostly roots and leaves, but fruits and flower buds are also used. Some of the medical properties can be related to phenolic compounds.

4. Capparis orientalis and C. • are grown from seeds or cuttings; other species can be grown successfully from seeds.

ACKNOWLEDGMENTS We thank ~he staff of the herbaria and associated libraries B1SH, C, E,

G, HUB, HUJ, JEPS, K, INN, MA, MARSSJ, OXE P, RNG, RSA, and US, for their help. We also thank Prof. De Frossard, the staff of AGRU- CAPERS (Spain) and URCIMAR (Morocco), and the reviewers and ed- itor of Economic Botany.

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