Capparis spinosa subsp. cordifolia (Lam.) Fici, 2015

Fici, Silvio, 2015, A taxonomic revision of the Capparis spinosa group (Capparaceae) from eastern Africa to Oceania, Phytotaxa 203 (1), pp. 24-36 : 28-30

2. Capparis spinosa subsp. cordifolia (Lam.) Fici , comb. et stat. nov.

Basionym: Capparis cordifolia Lamarck (1785: 609) . Type: MARIANA ISLANDS. Isles Mariannes , no date, Sonnerat s. n. (holotype

P). Capparis mariana Jacquin (1797: 57) . Capparis spinosa L. var. mariana (Jacq.) Schumann (1887: 201) . Lectotype (here designated):

[icon] “ Capparis mariana ” in Jacquin, Pl. Hort. Schoenbr.: t. 109. 1797. Capparis sandwichiana Candolle (1824: 245) . Type: HAWAII. In ins. Sandwich, no date, Menzies s. n. (holotype BM). Capparis baducca Blanco (1837: 438) , [non Linnaeus (1753: 504)]. [nom. illeg.]. Capparis sandwichiana Candolle var. zoharyi O. & I. Degener (1961: 369). Lectotype (designated by St. John 1965): HAWAII. Kauai,

Polihale, 22 November 1960, O. & I. Degener 27254 (lectotype NY, isolectotypes BISH, GZU, TI).

Shrub erect or prostrate; branches 1–2 (–4) m long, in most cases with greyish indumentum when young, early glabrescent. Stipular thorns usually wanting, rarely persisting and up to c. 0.5 mm long. Leaf-blade ovate, elliptic to suborbicular, c. (1.5–) 2.4–6 (–8) x (1.9–) 2.5–5.2 (–6.7) cm, more or less fleshy, rounded, truncate or nearly subcordate at base, rounded, obtuse or emarginate at apex, usually mucronulate; petiole 0.8–2 (–4) cm long. Flowers axillary, solitary; pedicel 3–7.5 cm long, glabrescent. Calyx markedly zygomorphic, with posterior sepal deeply saccate, (2.2–) 2.5–4 cm long, (1–) 1.3–2.5 cm deep, other sepals 2–2.6 x 0.8–2 cm. Petals white, obovate, suborbicular to rhombic, upper pair (2.2–) 3–5.5 x 2–4 cm, the lower pair (2–) 2.7–3.5 (–4.5) x 1.5–3.2 (–3.5) cm. Stamens (80–) 100–190, with filaments 3–6 (–7) cm long and anthers c. 3–3.5 mm long. Gynophore 5–7 cm long, slightly hairy near the base; ovary c. 5–8 x 1–2 mm. Fruit ellipsoid or oblong, c. 2.5–5 x 1.3–1.7 (–2.5) cm, ribbed. Seeds many, reniform, 2–4 mm long. ( Fig 2 ).

Distribution:— Indonesia, Papua New Guinea, Philippines, Palau, Northern Mariana Islands , Guam, Caroline Islands, Nauru, Solomon Islands, Vanuatu, New Caledonia, Fiji Islands, Tonga, Samoa, American Samoa, Cook Islands, Niue, French Polynesia, Pitcairn Islands, Hawaii.

Habitat:— Rocky shores, cliffs; on coral, limestone and volcanic rocks; up to c. 350 m altitude.

Phenology:— Flowering January–December. Noctiflorous (St. John 1965, Stone 1970).

Notes:— This subspecies includes all the populations of Capparis spinosa widespread in the Pacific area, Papua New Guinea and Indonesia, corresponding in its delimitation to C. spinosa var. mariana sensu Jacobs (1965) . In Indonesia it is recorded in Sumba, Semau, Timor and Leti, in Papua New Guinea in the Morobe District, New Ireland and New Britain. In the Pacific area subsp. cordifolia is widespread in coastal habitats from the Mariana Islands southwards to Vanuatu and New Caledonia and eastwards to French Polynesia, Pitcairn islands and Hawaii. No material was examined from the Marshall Islands, Kiribati, Tuvalu and Tokelau, where the occurrence of this taxon needs to be confirmed. The plant cultivated in the Philippines was described by Blanco (1837) as C. baducca . According to the same Author ( Blanco 1845) it was introduced in Parañaque (Luzon) from the Mariana Islands.

Based on the taxonomic isolation of C. spinosa from the other representatives of the genus in Malaysia and Australasia, Jacobs (1960) argued that this species was probably introduced in the Indo-Pacific for cultivation in Post-Columbian time. Differently Stone (1970) stated that the Pacific populations are undoubtedly native, based on their occurrence in communities of purely native elements restricted to certain habitats and their inability to persist to anthropic disturbance.

Adopting a narrower species concept, St. John (1965) recognized in the Pacific area two species, i. e. C. cordifolia widespread in the western, central and southern islands, and C. sandwichiana endemic to Hawaii. The distinction of these two taxa appears to be based on variable, frequently overlapping, vegetative and reproductive characters. Furthermore the cinereous-puberulent indumentum of young shoots, petioles and buds characterizing C. sandwichiana var. zoharyi - described from Kauai—is sporadically present in populations of other islands in the Pacific.

According to Jacobs (1965) another species included by St. John (1965) in the C. spinosa group, C. subscordata Spanoghe (1841: 166) , is a synonym of C. quiniflora Candolle (1824: 247) belonging to sect. Monostichocalyx Radlkofer (1884: 101).

With regard to C. mariana , in the protologue Jacquin (1797) included a detailed description of the plant cultivated in the Schönbrunn Garden, and the following note: “Crescit in insulis Marianis. In caldario floret Septembri & Octobri”. No original material of this taxon was found at W (A. Löckher, pers. comm.), nor was recorded by Savage (1945) at LINN, where a large amount of Jacquin items is present ( D’Arcy 1970). Thus, the only element of original material available for typification is the coloured plate included by Jacquin (1797, t. 109), which is here designated as lectotype ( Fig 3 ).

Uses:— Blanco (1837, 1845) reported in the Philippines the same use of capers as in Spain, while Jabobs (1960, 1965), recording the cultivation of the plant in Luzon, stated that “early governors of Guam exported pickled unripe capsules in considerable quantities”. In the latter island the present use of a relish, prepared by pickling the flower buds or sometimes the young fruits, is described by Stone (1970). According to St. John (1965) in the Hawaii the plant, locally named “pilo” or “maiapilo”, is valuable for the medicinal properties of its root, traditionally used for sprains, bruises, broken bones, as well as for fresh blending cuts. In herbarium labels of material from the Cook Islands the plant is said to be poisonous for goats.

Specimens examined:— INDONESIA. Sumba, near Waingapu near Kemenuru R., 06 April 1965, Kostermans & Wirawan 362 (K) ; INDONESIA. Coepang, Timor , June 1818, Cunningham 363 (K) ; INDONESIA. Timor , December 1898, Reeds s. n. (K) ; INDONESIA. Timor , no date, s. c. (G-DC) ; INDONESIA. Leti , August 1883, Riedel & Merper s. n. (K) ; PAPUA NEW GUINEA. Morobe District: Subdistrict Finschafen, Sialum, Henty & Katik NGF 49834 (K) ;

PHILIPPINES. Luzon , Rizal Province, Malabon, May 1914, Merril 516 (K) ; PALAU. Angaur Island, 02 January 1946, St. John 21500 ( BISH) ; PALAU. Angaur, 26 July 1946, Fosberg 25920 ( BISH) ; PALAU. Beliliou , 27 July 1946, Fosberg 26004 ( BISH) ; PALAU. Ngeruktabel Island , 17–19 March 1950, Fosberg 32198 ( BISH) ; PALAU. Olopshacal , 06 May 1936, Takamatsu 1475 (K) ; NORTHERN MARIANA ISLANDS . Farallon de Medinilla, 04 March 1984, Herbst 7516 ( BISH) ; NORTHERN MARIANA ISLANDS . Maug, West Island , 25 February 1984, Herbst 7373 ( BISH) ; NORTHERN MARIANA ISLANDS . Tinian, July 1933, Kanchira 2277 (K) ; GUAM. Cocos Island , 24 June 1962, Stone 4240 (K) ; GUAM. Above Anao point, 01 August 1954, Moran 4544 (K) ; FEDERATED STATES OF MICRONESIA. Fais Island , 30 July 1965, Evans 359 (K) ; NAURU. s. l., 11 December 1990, Cherrier 45234 ( BISH) ; NAURU. Anabar, 28 November 1978, Fosberg 58636 ( BISH) ; NAURU. Menen , 03 January 1934, Burges K 17 (K) ; SOLOMON ISLANDS. Bellona , 06 May 1968, Sirute’e 9618 (K) ; SOLOMON ISLANDS. Bellona , 30 June 1933, Templeton Crocker Exped. s.n. ( BISH) ; SOLOMON ISLANDS. Santa Ana Island , 24 November 1969, Mauriasi 17218 (K) ; VANUATU. Malekula, Tisvel , 28 September 1971, Gillison RSNH 3547 (K, P) ; VANUATU. Loh Island , 01 November 1993, Curry 1403 (K) ; NEW CALEDONIA. s. l., s. d., Balansa 1697 (K) ; NEW CALEDONIA. Wallis, Lake Lalolalo , 11 November 2007, Pillon 997, Gatimel, Jourdan, Meyer & Richling (P) ; FIJI ISLANDS. Viti Levu, Nandronga, Thuvu , near Singatoka , 24 May 1942, Greenwood 917 (K) ; FIJI ISLANDS. July 1929, Vatu Lele I., Tothill & Payne s. n. (K) ; FIJI ISLANDS. Kadavu, near Cape Washington , 28 October 1933, Smith 319 (K, BISH) ; FIJI ISLANDS. Lau Group, Aiwa , 30 August 1924, Bryan 494A ( BISH) ; FIJI ISLANDS. Rotuma Group, Solkope, 24 August 1938, St. John 19702 ( BISH) ; TONGA. Tongatapu, Hufangalupe , 01 June 1926, Setchell 15629 ( BISH) ; TONGA. Tongatapu, Fangaveha , 07 June 1953, Yuncker 16250 ( BISH) ; SAMOA. Savai’i, Falelima , May 1907, Vaupel 630 (K) ; SAMOA. Savai’i, Falealupo , 29 September 1931, Cristophersen 2782 (K) ; SAMOA. Savai’i , 21 October 1973, Wistler 995 (K) ; AMERICAN SAMOA. Manu`a Islands , Ta`u, 22 August 1975, Whistler W 3222 (K, BISH) ; COOK ISLANDS. Aitutaki , 03 September1969, Stoddart 2297 ( BISH) ; COOK ISLANDS. Mitiaro , 26 April 1985, Whistler 5621 ( BISH) ; COOK ISLANDS. Mitiaro , 02 July 1991, Luttrell s. n. (K) ; COOK ISLANDS. Akaimi Island, Aitutaki , 19 August 1955, Mc Kee 3025 (K) ; COOK ISLANDS. Rarotonga, Motu Tapu , 15 August 1929, Wilder 840 ( BISH) ; COOK ISLANDS. Rarotonga, Ngatangiia Harbour , July 1899, Cheesman 506 (K) ; NIUE. Makefu , 07 February 1940, Yuncker 10112 (P) ; FRENCH POLYNESIA. Rurutu , 20 April 1981, Hallè 7141 (K, P) ; FRENCH POLYNESIA. Rurutu , 22 April 1981, Nicolas Hallè 7227 (P) ; FRENCH POLYNESIA. Rurutu, Arapiia, 02 September 1934, St. John 16763 & Cooke (P) ; FRENCH POLYNESIA. Rimatara, Anapoto, 04 September 1934, St.John 16850 ( BISH) ; FRENCH POLYNESIA. Rurutu, Mato Arei , 10 February 1921, Stokes 59 ( BISH) ; FRENCH POLYNESIA. Austral Islands, Rapa, Vavai , between Puero Bay and Kuturoa Pt., 11 April 2002, Perlman 18052, Fenstemacher & Motley (P) ; FRENCH POLYNESIA. Tuamotu, Ile Makatea , 29 April 1982, Florence 3034 (K) ; FRENCH POLYNESIA. Tuamotu Ilsands, Makatea , 17 September 1932, Wilder 1117 (K) ; PITCAIRN ISLANDS. Pitcairn I., no date, Withney Expedition 107 ( BISH) ; PITCAIRN ISLANDS. Henderson I., 07 May 1991, Florence & Waldren 10851 (K) ; PITCAIRN ISLANDS. Henderson I., 15 May 1991, Florence, Chepstov-Lusty & Waldren 10912 ( BISH, K, P) ; PITCAIRN ISLANDS. Henderson , 3–4 February 1957, Lintott H. 24 ( BISH) ; HAWAII. Kealakekua , 11 March 1950, Greenwell &. Nateway s.n. (K) ; HAWAII. s. l., 1879, Gaudiebaud 262 (K); HAWAII. Pearl & Hermes Reef, N. Island , 18 May 1923, Wilder 2 ( BISH) ; HAWAII. N. Kona, Kahului , 28 May 1948, Alexander & Kollogg 5364 ( BISH) ; HAWAII. Lanai, Limestone Point , 31 Mar 1914, Munro s.n. ( BISH) ; HAWAII. Kauai, Polihale , 27 November 1960, O. & I. Degener 27256 ( FI) ; HAWAII. Kauai, Polihale Beach , 24 Sep 1977, Corn s.n. ( BISH) ; HAWAII. Oahu , 7 May 1932, Larsen s.n. ( BISH) ; HAWAII. Oahu, Kailua, Popoi`a Island , 8 Nov 1932, Fosberg 8891 ( BISH) ; HAWAII. Oahu, Pounders Beach , 9 August 1987, Perlman 6280 ( BISH) ; HAWAII. E. Maui, Kahikinui , November 1910, Rock 8694 ( BISH) ; HAWAII. s. l., 15 July 1968, O. & I. Degener 31657 ( FI, TI) .

Figures

FIGURE 2. Capparis spinosa subsp. cordifolia. A. Flower. B. Ripe fruit. C. Plant in natural habitat (Hawaii). (Photos A, C by G. Carr, B by M. LeGrande).

FIGURE 3. Lectotype of Capparis mariana (from Jacquin 1797).