Capparis spinosa subsp. cartilaginea (Decne.) Maire & Weiller

Fici, Silvio, 2015, A taxonomic revision of the Capparis spinosa group (Capparaceae) from eastern Africa to Oceania, Phytotaxa 203 (1), pp. 24-36 : 25-27

1. Capparis spinosa subsp. cartilaginea (Decne.) Maire & Weiller in Maire (1965: 120).

Capparis cartilaginea Decaisne (1835: 273) . Lectotype (designated by Rivera et al. 2003): EGYPT. Lassaf Arab, Désert du Sinai, Juin 1832, Bové 148 (lectotype P, isolectotypes K, L).

Capparis inermis Forsskål (1775: 100) [non Turra (1780: 65)]. Neotype (designated by Rivera et al. 2003): YEMEN: Aden: Jebel Shamsan, Tower of Silence and vicinity, 7 June 1987, Boulos, Gifri, Hussein & Awad 15560 (neotype K).

C apparis galeata Fresenius (1836: 111) ; Capparis spinosa L. var. galeata (Fresen.) Hooker f. & Thomson (1872: 173) Type: ETHIOPIA. Abyssinien, Halei, no date, Rüppell s. n. (holotype FR, lost). Neotype (here designated): ETHIOPIA. Harar Province: near Argoba Village west of Bisidimo, about 15 km. SSW of Harar, 07 September 1963, Burger 3227 (neotype K, isoneotype B).

Capparis pyracantha Bojer (1843: 59) ; Capparis spinosa L. var. pyracantha (Bojer) Hadj-Moustapha (1965: 16) . Type: MADAGASCAR. “Patria: insula Madagascar, crescit in arenosis maritimis, sinu Saint-Augustin orâ austro-occid.” Bojer s. n. (holotype W or P, lost). Neotype (here designated): MADAGASCAR. Track from Toliara to Fanantsony, Saint Augustin, 06 August 1993, Lewis, Du Puy, Hardy & Rapanarivo 533 (neotype P).

Capparis uncinata Edgeworth (1847: 1213) . Type: YEMEN. Aden, 1846, Edgeworth s. n. (holotype, not known, presumed lost). Neotype (here designated): YEMEN. Aden, February 1837, Thomson s. n. (neotype K).

Capparis antanossarum Baillon (1885: 462) . Type: MADAGASCAR. Pays des Antanosses émigrés et forêt de Lavanala, October 1869, A. Grandidier s. n. (holotype P).

Capparis galeata Fresenius var. montana Schweinfurth (1896: 191) . Type: YEMEN. Menacha, 1889, Schweinfurth 1920 (holotype B).

Capparis galeata Fresenius var. lanceolata Blatter & Hallberg (1918: 724) . Type : IRAN. Persian Baluchistan: Sib., 18–24 July 1916, Hotson s.n. (syntype BLAT); IRAN. Gusht, N. W. of Dizak , August 1916, Hotson s.n. (syntype BLAT) .

Shrub with branches mostly erect, sometimes spreading or scrambling, 0.6–4 m tall; twigs firstly white-pubescent, early glabrescent. Stipular thorns mostly vigorous, recurved, 2–4 mm long, yellowish, sometimes wanting. Leaves more or less fleshy, when dried subcoriaceous, blade ovate, elliptic or orbicular, glabrous, (2–) 2.5–6.2 x (1.5–) 2–5.5 (–6) cm, rounded, cordate or acute at base, rounded, emarginate to acute at apex, mostly with recurved mucro up to c. 1.5 mm long; petiole (0.7–) 0.9–2.7 cm long. Flowers solitary, axillary in the upper part of twigs, strongly zygomorphic, noctiflorous; pedicels (3.5–) 4.5–9.5 (–12) cm long. Calyx with posterior sepal deeply saccate, 2.5–4.5 cm long, 1.7– 2.5 cm deep, the others concave, boat–shaped, (1.5–) 2–2.5 x 1–1.3 cm. Petals white, upper pair connate, lower pair free, obovate or obdeltoid, c. 3–5 x 2.5–3 cm. Stamens up to c. 200, with filaments c. 3–5 cm long and anthers 2–3 x 0.6–0.7 mm. Gynophore (3–) 3.5–5 cm long in flower, glabrous; ovary oblong-ellipsoid, c. 5–6 x 2 mm. Fruit borne on a thickened stalk, ovoid or ellipsoid, 3–5 (–9) x 2–3 (–4) cm, ribbed, usually reddish, many-seeded. ( Fig. 1 ).

Distribution:— Egypt, Sudan, Chad, Ethiopia, Eritrea, Djibouti, Somalia, Kenya, Tanzania, Seychelles, Madagascar, Yemen, Oman, Saudi Arabia, Israel, Jordan, Iraq, Iran, Pakistan, India. This taxon has also been recorded in the Mascarene Islands, Kuwait and Afghanistan ( Zohary 1960, Elffers et al. 1964, Jacobs 1965).

The occurrence in southern Iraq was reported by Rechinger (1964) for the area south of Al-Busaiyah. In north-western India this taxon was recorded in the Kutch District by Blatter (1908), and at Dwarka in the Kathiawar Peninsula by Jacobs (1965).

Habitat:— Cliffs, rocky slopes, stony wadis, sandy and alluvial soils; on limestone, sandstone, gypsum, coral, gneiss and volcanic substrata. From sea level to c. 2400 m.

Phenology:— Flowering January-December. In herbarium labels the plant is frequently described as noctiflorous; in Somalia the flower is recorded as opening c. 19.30 and fading c. 11.00 (Gillett et al. 22135).

Notes:— According to Jacobs (1965) the differential characters of this taxon, including the posterior sepal deeply saccate, are only quantitative, suggesting that it would not deserve specific rank besides Capparis spinosa . Due to its peculiar features and distribution, it is here treated at subspecific rank within C. spinosa , as proposed by Maire & Weiller in Maire (1965). On the contrary Ellfers et al. (1964) treated this taxon at specific rank - sub C. cartilaginea - yet underlining that it is closely related to C. spinosa var. spinosa , differing in the flower markedly zygomorphic and in the recurved mucro at the leaf apex. According to the same authors C. cartilaginea is also similar to the form widespread in several islands of the Pacific Ocean [sub Capparis spinosa L. var. mariana (Jacq.) Schumann (1887: 201) ], which is differentiated by its orbicular leaves without mucro. Also Inocencio et al. (2006) treated this taxon at specific rank, but included it in a different section, sect. Galeatae Inocencio et al. (2006: 126), on the basis of the flowers always solitary at leaf axils and the abaxial sepal strongly galeate.

If compared with other representatives of the group, especially the Mediterranean ones, subsp. cartilaginea is characterized by the habit usually erect and the fleshy leaves, testifying its closeness with the primitive xerotropical stock of this complex. Zohary (1960) regarded this taxon in the Near Eastern countries as a relic of a tropical flora which dominated the region during the Late Tertiary. Furthermore, according to Hedge & Lamond (1970) it is an example, not infrequent in the Capparaceae , of a “Sudanian element” occurring in tropical East Africa and also in the Arabic Peninsula and Baluchistan. In the Horn of Africa it is widespread in deciduous, evergreen and semidesert open bushland on various substrata ( Fici 1993).

Several taxa described in the past from eastern Africa, Madagascar and southern Asia on the basis of quantitative characters, fall within the variability of this subspecies. Lobin (1999) stated that the type specimen of C. galeata , collected in Abyssinia by Rüppell, is missing at FR, and underlined that it was not mentioned in the catalogue furnished by Geyler (1867 –87). The absence of original material in FR was confirmed during the present research by S. Dressler (pers. comm.); a neotype is therefore designated. Also for Capparis pyracantha the holotype, collected by Bojer in Madagascar, is missing at either W and P (A. Löckher and J. Florence, pers. comm.), and a neotype is here designated.

With regard to C. uncinata the type specimen, collected by Edgeworth at Aden in 1846, was not found at K, OXF and G (J. Osborne, S. Harris and L. Gautier, pers. comm.) and it has not been seen at other herbaria in the present study. Being the original material presumably lost, a neotype is here designated.

In northern Africa, Maire (1965) recorded C. spinosa subsp. cartilaginea in the “Sahara septentrional tripolitain: a l’Est de Ghadamès”, based on material collected by Largeau (P). According to Fici (2014) this material belongs to subsp. rupestris (Sm.) Nyman (1878: 68) , as probably other records of subsp. cartilaginea from Tunisia ed Algeria ( Jafri 1977).

Uses:— In eastern Africa the plant is frequently utilized for its properties. The following uses are reported in herbarium labels. In Somalia the leaves steeped in hot water are utilized as a treatment of cough in adults, children and even infants (Peck 226, Aronson et al. 34). The same use is recorded for the fruit in the northern part of the country, where the native name “Goh” represents the sound of a cough (Collenette 24). In this area the fruit is eaten, and the dried leaves mixed with wood ash are used as “chewing tobacco”, while the plant is grazed by all stock, except cows. Infusions of stems and leaves are used in Kenya to cure animals which are sick after giving birth (Matthew 6669).

Specimens examined:— EGYPT. Between Kosser and Ras-Benas , 29 June1864, Schweinfurth s.n. (K) ; EGYPT. Wadi Kalfa , February 1906, Mushler s.n. (K) ; EGYPT. Wadi Umm Grifat, 68 km north of Mersa Alam , 13 March 1964, Boulos s.n. (K) ; EGYPT. Sinai, Wadi El Humur , 14 April 1937, Shabetai 210.520 (K) ; EGYPT. Sinai, St. Katerine region, Wadi Zaghra , 27 Novemberr 1999, Boulos & Ali s.n. (K) ; EGYPT. Sinai, between Ras Sudr and Abu Zneima , 17 April 2003, Boulos s.n. (K) ; SUDAN. Red Sea, Ruwaya , January 1922, Johnston 214 (K) ; SUDAN. Nubia about 21°, Int. sea coast, 1896, Bent s.n. (K) ; SUDAN. Soturba-Gebirges, Gebel Hotarger , 30 April 1864, Schweinfurth s. n. (K) ; SUDAN. Nubische Küste, Insel Macaus , 20 June 1864, Schweinfurth s.n. (K, P) ; CHAD. Ennedi, Archei , 10 September 1935, Dupont & Murat 337 (P) ; ETHIOPIA. c. 10 km SE of Diredawa , 01 February 1966, de Wilde & de Wilde-Duyfjess 9878 (K) ; ETHIOPIA. Shewa, 2 km west of Metahara , 20 May 1974, Gilbert 3461 (K) ; ETHIOPIA. El Sod , 15 November 1952, Gillett 14272 (K) ; ETHIOPIA. Sidamo Region, Mt. Furolli , 17 December 1997, Friis, Bidgod, Legesse, Wandefrash & Gibre-Iwoth 8673 (K) ; ETHIOPIA. Shoa Province , south side of Blue Nile gorge, 22 May 171, Ash 864 (K) ; ETHIOPIA. Abyssinie , 01 June 1860, Russel s. n. (P) ; ETHIOPIA. Between Nazareth and Awash-Station , 27 January 1966, De Wilde 9808 (P, B) ; ERITREA. Ile Dalak , s. d., Dillon s. n. (P) ; SOMALIA. 3 mls E of El Dad , 26.09.1956, Bally B10859 (G) ; SOMALIA. Bari Region: Dud-Shabel, 28 km from Bosaso on road to Karin , 04 December 1985, Thulin & Warfà 5618 (K) ; SOMALIA. Las Anod , 13 August 1941, Peck 38 (K) ; SOMALIA. Erigavo, Burran , 07 October 1941, Peck 226 (K) ; SOMALIA. 21 km S of Garowe , 14 July 1980, Gillett 23144 (K) ; SOMALIA. Baran , 20 October 1983, Aronson, Ikar, Linington, Wedderburn & Wickens 34 (K) ; SOMALIA. Buran, Sorl , 06 September 1929, Collenette 34 (K) ; SOMALIA. Garoe , 06 August 1969, Lanza 25 ( FT) ; SOMALIA. Oltregiuba, Sar Uanle , 09/ 11 June 1973, Bavazzano s. n. ( FT) ; SOMALIA. 05°57’N – 48°57’E, 28 May 1979, Gillett, Hemming & Watson 22135 (K) ; DJIBOUTI. Vallèe d’Assamo, au SE d’Ali Sabreli , 30 April 1938, de la Rüe s. n. (P) ; DJIBOUTI. Yager , 16 april 1986, Cesar 2378 (P) ; KENYA. Turkana District: Lokapilak , 05 August 1968, Mwangangi & Gwinne 1071 (K) ; KENYA. Turkana District: South Turkana, near Mugurr , 06 June 1970, Matthew 6669 (K) ; KENYA. Mombasa District: Mombasa, February 1876, Hildebrandt 1922 (K) ; KENYA. Mombasa District: Malindi , October 1951, Twredie 970 (K) ; KENYA. Baringo District: Lake Bogoria , 03 April 1977, Hoopper & Townsend 1627 (K) ; KENYA. Lamu District: Patta, I. Lamu, December 1967, Kirk 9/68 (K) ; KENYA. Kilifi District: Vipingo, Kijangurani , 02 January 1974, Frazier 845 (K; EA) ; KENYA. Marsabit District: Laisarnis , July 1933, Gardner s. n. ( MA) ; KENYA. Marsabit District: Nalawaru , 12 February 1953, Gillett 15125 (B) ; SEYCHELLES. Aldabra Group, Middle Island , 28 January 1968, Fosberg 49090 (K) ; SEYCHELLES. Aldabra Group, Polymnie Island , 26 August. 1967, Stoddart 997 (K) ; SEYCHELLES. Aldabra Group, Astove Atoll , 06 October 1967, Gwynne & Wood 1291 (K) ; SEYCHELLES. Aldabra Group, West Island , 09 January 1968, Renvoize 720 (K) ; SEYCHELLES. Aldabra Group, South Island , 19 January 1968, Fosberg 48931 (K) ; MADAGASCAR. Tulear, Manombo, Mikea Forest , 05 December 1993, Labat, Du Puy & Comtet 2457 (K) ; MADAGASCAR. Tulear Province, 35 km southeast of Tulear near Onilahy River , 06 December 2000, Rakotondranony, Razafimanonjy & Smith 69 (K) ; MADAGASCAR. Antsiranana, Ambilobe, Matsaborimanga, Ankarana Reserve , 18 November 1997, Bardot-Vaucoulon & Andrianantoanina 970 (K, P) ; MADAGASCAR. Near Tulear , s.d., Richey 7 (K) ; MADAGASCAR. Vallée du Fiherenana , November 1933, Humbert 11579 (P) ; MADAGASCAR. Vallée de l’Onilaky , 05 February 1947, Humbert 20142 (P) ; MADAGASCAR. Estuarie de l’Onilahy , côté sud, 20–21 March 1955, Humbert 29529 & Capuron (P) ; MADAGASCAR. Lac Eotry , 11 August 1898, Grandidier s. n. (P) ; YEMEN. Aden, Jebel Shamsan , 07 June 1987, Boulos, Gifri, Hussein & Awad 16560 (K) ; YEMEN. Aden, 24 April 1876, de Marchesetti (K) ; YEMEN. Aden, 1885, Chaper s. n. (P) ; YEMEN. Aden, 26 November 1888, Schweinfurth s. n. (P) ; YEMEN. Uossil , 08 February 1889, Schweinfurth 1265 (K) ; YEMEN. Mountains behind Dafrd , 26 August 1982, Bisset 160 (K) ; YEMEN. Attara , 14 May 1887, Deflers 410 (P) ; YEMEN. Socotra , October 1880, Balfour 192 (K) ; YEMEN. Socotra, J. Umhari , 18/ 25 October 1966, Virzo S 32 (K) ; YEMEN. Socotra, Hammaderoh , 06 April 1967, Smith & Lavranos 254 (K) ; OMAN. Remacs Wadi , 21 June 1973, Mc Lead 3 (K) ; OMAN. Mina Al Fahal , 17 September 1979, Miller & Whitcomb 2014 (K) ; SAUDI ARABIA. Yanbu ‘al Bar , 1972, Collenette s. n. (K) ; SAUDI ARABIA. Makna Range, Jebel Al Hamtha , 05 May 1978, Collenette 728 (K) ; SAUDI ARABIA. Al Aflaj – Riyadh, 12 October 1966, Mosnier 2697 (P) ; ISRAEL. Wadi Klabis , 21 January 1963, Davis 3069 (K) ; JORDAN. Aqaba, Wadi Rum , 22 September 2004, Abulaila & Tehabsem 2004JOR323-2 (K) ; IRAN. Fars, Tarq-i-Khar , 13 May 1959, …. 3659 (K) ; PAKISTAN. Beluchistan , coast, Pierce 480 (K) ; PAKISTAN. Sind, s.d., Dalzell s.n. (K) ; PAKISTAN. Karachi, Kunashu , October 1847, Ellerton Stocks 561 (K) .

Figures

FIGURE 1. Capparis spinosa subsp. cartilaginea. A. Flower. B. Opening flower showing the deeply saccate posterior sepal. C. Plant in natural habitat (Sinai). (Photo A by S. Cambria, B by F. M. Raimondo).