Research Article |
Corresponding author: Alexander P. Sukhorukov ( suchor@mail.ru ) Academic editor: Gianpietro Giusso del Galdo
© 2016 Alexander P. Sukhorukov, Maria Kushunina.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Sukhorukov AP, Kushunina M (2016) Taxonomic revision and distribution of herbaceous Paramollugo (Molluginaceae) in the Eastern Hemisphere. PhytoKeys 73: 93-116. https://doi.org/10.3897/phytokeys.73.10365
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The genus Paramollugo with the type species Paramollugo nudicaulis (≡Mollugo nudicaulis) has recently been described after molecular investigations. Here we report two new endemic Malagasy species: Paramollugo simulans and P. elliotii, and transfer a forgotten New Caledonian endemic Mollugo digyna to Paramollugo (P. digyna). Consequently, the number of Paramollugo species in the Eastern Hemisphere is increased from three to six. Almost all genus representatives (except P. nudicaulis, which has a wide distribution in Southern Asia, Arabia and tropical Africa) are endemic to Madagascar, Somalia, or New Caledonia. Since the type of seed coat ornamentation is crucial for species delimitation, a diagnostic key with new taxonomically significant carpological characters and other new traits is provided for all the herbaceous Paramollugo. The distribution patterns of P. nudicaulis s.str., P. simulans and P. elliotii are presented.
Madagascar, Mollugo , New Caledonia, new species, seed characters, taxonomy
The new genus Paramollugo Thulin has recently been split from Mollugo s.l. after molecular investigations (
We have previously shown that seed morphology and ultrasculpture are very useful for the diagnostics and taxonomy of major groups within the former Mollugo s.l., and that the ‘Mollugo nudicaulis’ group (=Paramollugo) needs further taxonomic studies (
The revision of herbarium material was undertaken in the collections B,
The seed surface of Paramollugo was examined using scanning electron microscopy (SEM; JSM–6380, JEOL Ltd., Japan) at 15 kV after sputtercoating with gold-palladium. Fallen fruits (1–2) from the following vouchers were used for SEM:
Paramollugo digyna (Montrouz.) Sukhor.: NEW CALEDONIA. Canala, [without date] anonym 236 (
P. elliotii Sukhor.: MADAGASCAR. Ambongo, 14 February 1841, M. Pervillé 643 (P04582808); Mailake prov., February 1890, M. Douillot s.n. (P04582888); Baie de Baly, January 1905, H. Perrier de la Bathie 9211 (P04582813); Majunga, 27 April 1912, Afzelius s.n. (K); Environs de Majunga, December 1924, H. Humbert & H. Perrier de la Bathie 4020 (P04582873);
P. nudicaulis (Lam.) Thulin: BHUTAN. Tashigang distr., Dangme Chu valley, 3000 ft, 17 August 1915, Cooper & Bulley 4498 (
P. simulans Sukhor.: MADAGASCAR. Vaingaindrano, [without date] Scott Elliot 2257 (
Six Paramollugo species in the Eastern Hemisphere (P. angustifolia, P. decandra, P. digyna, P. elliotii, P. nudicaulis, P. simulans) are accepted in the present paper, instead of the three previously recognized species (P. angustifolia, P. decandra, and P. nudicaulis). The core species, P. nudicaulis s.str., is found to be widely distributed in the tropics of the Eastern Hemisphere as a weed species (see Fig.
The genus Paramollugo in the Eastern Hemisphere is one of the most morphologically difficult groups in Molluginaceae. We found that the following reproductive characters are useful for delimiting the Paramollugo species: size of the perianth segments, length of the anthers, number of stylodia and locules in the capsule, and seed shape and ornamentation. Seed surface is either papillate (P. digyna, P. decandra, P. nudicaulis; Fig.
Paramollugo decandra is not included in the text; it is a small shrub (this character differentiates this species from all other Paramollugo in this region, which are herbs)
1 | Seeds with well-visible papillate ornamentation and warty or striate outgrowths of the testa cells | 2 |
– | Seeds without distinct papillae | 3 |
2 | Annual; leaves obovate or oblong; capsules three-valved; outer wall of the testa cells striate, without warty outgrowths | P. nudicaulis (tropical Africa and Asia) |
– | Annual or perennial herb; leaves reniform or almost triangular; capsules usually two-valved; outer wall of the testa cells with several or numerous warty outgrowths | P. digyna (New Caledonia) |
3(1) | Perennial herb with numerous (25–50) linear or lanceolate sessile leaves to 5 mm wide, not divided into petiole and lamina; anthers 0.55–0.70 mm; outer perianth segments 2.6–3.0 mm | P. elliotii (Madagascar) |
– | Annuals with 5–20 leaves of different shape; anthers 0.25–0.35 mm; perianth segments at fruiting stage not exceeding 2.5 mm in length | 4 |
4 | Leaves linear, to 2 mm wide; seeds 0.35–0.45 mm in diameter | P. angustifolia (Somalia) |
– | Leaves usually oblong to obovate, to 12 mm; seeds 0.55–0.7 mm in diameter | P. simulans (Madagascar) |
MADAGASCAR, [Atsimo-Andrefana region], Toliara [district], Belalanda, Ranobe, 4 km a l’Est du village Andrevo, 22°58'20"S, 43°36'56"E, 135 m, 17 March 2006, R. Ranaivojaoana et al. 1442 (holotype K000607662; isotype BR0000000514054, P05196698). (Fig.
Paramollugo simulans is morphologically similar to P. nudicaulis but differs by having colliculate seed surface (testa cells slightly convex, not papillae-like) with no striate secondary cell ornamentation. Some forms of P. simulans with linear leaves look like P. angustifolia, but they reliably differ by the seed size (0.55–0.7 mm and 0.35–0.45 mm, respectively).
Annual to 25(30) cm; leaves (5–9)10–25, rosulate, rather thin, obovate, narrowly oblong, sometimes lanceolate, 20–50 × 2–12 mm, continuously tapered in petiole (up to 20 mm) or rarely not clearly divided into petiole and blade if lanceolate or linear, glabrous or shortly papillate; stems many (rarely 1 in underdeveloped exemplars), branched in the middle part with many lateral branches forming loose inflorescence with semi-spherical habit, always with short (up to 20 mm) sterile branches (not carrying flowers); peduncles to 15 mm, basally with white or brownish bract to 1.5 mm; flowers 3–4 mm in diameter when open; perianth segments 1.3–1.7 mm long in flowering, 2–2.4 mm long in fruiting, ovoid, green from abaxial side, white or pinkish inside; stamens 5 to 10, anthers elliptic, 0.30–0.35 mm; capsule 3-valved, ovoid, 1.5–2.3 mm, contains (8)10–18 seeds; seeds are reniform, dark red or almost black, 0.55–0.7 × 0.6 × 0.3 mm, with colliculate ultrasculpture.
Sandy, rocky, ruderal sites or dry forests at elevations 0–1700 m.
November–June.
MADAGASCAR: Alaotra-Mangoro region: Moramanga, Andasibe, Berano, Ambatovy forest, Analamay, 18°49'05"S, 48°19'25"E, 1145 m, 24 February 2005, P. Antilahimena et al. 3464 (P05196697); Androy region: Beloha, 25°05'S, 45°10'E, alt. 125 m, deciduous thicket with Indigofera & Crotalaria, locally common, 1 December 1989, Dupuy 2448 (K); Anosy region: Fort Dauphin [Tolanaro], St. Luce, 1 May 1989, N. Dumetz 720 (P04582891); Atsinanana region: Vaingaindrano, [without date] Scott Elliot 2257 (
Although a rather limited number of specimens is available, it is clear that P. simulans can occupy disturbed habitats and is therefore unlikely to be at risk of extinction. Following the
The specific epithet “simulans” means “imitating” and refers to the morphological similarities of the new species and Mollugo nudicaulis.
P. elliotii is morphologically similar to both P. nudicaulis and P. simulans, but differs by perennial life form, narrow (linear to oblanceolate), (sub)sessile leaves, larger anthers (0.55–0.70 mm), and larger outer perianth segments (2.6–3 mm) at the fruiting stage (P. nudicaulis and P. simulans are annuals, with broader, usually petiolate leaves, anthers of 0.25–0.30 mm and smaller perianth to 2.4 mm long).
Perennial herb to 35–40 cm; leaves 25–50, rosulate, thick, linear to oblanceolate, (10)20–60 × 0.5–5.0 mm, sessile or not clearly divided into petiole and lamina, glabrous; stems several, stout, branched in the middle part and forming loose semi-spherical inflorescence with numerous lateral branches, sterile (up to 30 mm) branches always present; peduncles to 15 mm long, basally with white or brownish bract 0.7–1.2 mm long; opened flowers 3.5–4.5 mm in diameter; perianth segments ovoid, green on the abaxial side, white or pink inside, 1.5–2.0 mm in flowering, unequally enlarging at fruiting stage (three outer segments 2.6–3.0 mm long, two inner 2.3–2.6 mm long); stamens 5, anthers elliptic, 0.55–0.7 mm long; capsule 3-valved, ovoid, ca. 2 mm long, with (9–14)15–21 seeds; seeds reniform, black, 0.55–0.7 × 0.5 × 0.3 mm, with colliculate surface.
All examined specimens were collected on the sand dunes near the sea at lower elevations 0–300 m. We assume that P. elliotii is a psammophilous species.
All examined specimens (collected in January, February, July, September or December) bear both flowers and fruits. However, the exact phenology has to be investigated more precisely, since it can depend on the geographical latitude and other conditions.
MADAGASCAR: Atsimo-Atsinanana region: Ambongo, 14 February 1841, M. Pervillé 643 (K, P04582808; P04582809; P04582810); Melaky region: province de Mailake, Namela, dunes, February 1892, M. Douillot s.n. (P04582869, P04582886); Baie de Baly, January 1905, H. Perrier de la Bathie 9211 (P04582813; P04582814); along Manambolo river, 19°09'S, 44°49'E, 50 m, 2 December 1996, C.C.H. Jongkind 3342 (BR0000000555871); Boeny region: Majunga, 27 April 1912, Afzelius s.n. (K); Majunga, dunes, 24 December 1920, H. Poisson 44 (P04582831); Environs de Majunga, dunes, December 1924, H. Humbert & H. Perrier de la Bathie 4020 (K, P04582873) & 2020 (P04582874; P04582877);
Endemic to Madagascar (Fig.
All except one specimen are nearly a hundred years old or even older, and no information is available on the current extent of the wild populations of this species. Therefore P. elliotii is given a Data Deficient (DD) status (
The specific epithet is given after after George Francis Scott Elliot (1862–1934), a British botanist who provided significant contributions to the flora of Madagascar.
Basionym: Mollugo digyna Montrouz., Ann. Acad. Roy. Sci. Lyon, Sect. Sci. 10: 179 (1860).
(Sukhorukov, selected here). NEW CALEDONIA. West face of Massif de Koniambo, 300 m, iron-serpentine scrub on track through “maquis”, 11 October 1963, P.S. Green 1287 (K001045648!).
This taxon described by X. Montrouzier was completely forgotten and cited as Mollugo nudicaulis in the treatments of the New Caledonian flora (e.g.,
No original material of Paramollugo digyna was traced. The herbarium of X. Montrouzier, who described M. digyna, is located in University of Montpellier herbarium (
Paramollugo digyna is known only from New Caledonia where it usually grows on rocky or stony substrates; locally common (
(selected specimens). NEW CALEDONIA: Kanala [Canala], 1859–1860, Viellard (
We thank Caroline Loup, curator of herbarium
Specimens of Paramollugo nudicaulis from the Eastern Hemisphere and P. angustifolia examined in the herbaria visited.
P. angustifolia:
SOMALIA, Bay region, c. 3 km SW of Dilinsoor, near Buur Dilinson, 2°24'N, 42°58'E, 20 May 1990, Thulin, Hedrén & Abdi Dahir 7606 (K!, B! – isotypes).
Paramollugo nudicaulis:
ASIA:
BHUTAN: Tashigang distr., Dangme Chu valley, 3000 ft, 17 August 1915, Cooper & Bulley 4498 (
CEYLON: Bolawatta, 16 July 1932, Simpson 9864 (
CHINA: Hainan: Ngai distr., Naam Shan Leng, 17 July 1932, S.K. Lau 271 (P04601248, E); Ch’ang-kiang [Changjiang] distr., 23 May 1933, Lau 1788 (
INDIA: nr Madras [Chennai], 1856, Cleghorn s.n. (E); Madras [presidency], October 1885, Gamble 18217 (
MYANMAR: Upper Burma [without precise location] 12 September 1990, Huk s.n. (E).
PAKISTAN: Khyber Pakhtunkhwa province: Akora, 1862, Griffith 2471 (K); Mansehra, 29 September 1888, Duthie 7480 (K); Swat state [district], nr Madian [Madyan], 14 August 1952, Rodin 5508 (K); Swat, Najigram, 3 August 1953, Stewart & Rahman 25421 (
SAUDI ARABIA: [Jizan region] nr Ad Darb, Abha-Gijan road, 1 November 1985, Collenette 5444 (E); [Al Bahah region] Bani Sharfa, 27 November 1966, M. Mosnier 2922 (P04934292).
VIETNAM: [Binh Thuan province] Phan-Rang prov., Cana, 24 December 1923, Poilane 9318 (K); Quang Ngai province, Dong Phu, June 1934, Petelot 5347 (P04601246, P04601249); [Ninh Thuan province] Phan Rang, 15 July 1930, Poilane 17872 (P04601247).
YEMEN: Socotra, 4 km SE of Hadiboh, 19 February 1989, Miller et al. 8225 (E); Sokotra, Hadiboh Plain, 24 January 1990, Miller et al. 10005 (E); [Al Hudaydah Governorate] Hais [Hays], 1837, M. Botta s.n. (P04601259); between Wusal al Ali & Beni Muftah, 17 September 1978, Wood 2476 (E); [Lahij Governorate] Wadi Labdah, nr At Tur, 17 November 1982, King 151 (E).
AFRICA:
ANGOLA: Benguella province: Membassoco, February 1942, Faulkner A131 (
BENIN: without precise locality, 1904, Forluo s.n. (P04583546); between Paouignan and Savalou, January 1902, M. Poisson s.n. (P04583577); without precise locality, 1902, M. Poisson s.n. (P04583579); near Agouagon, 24 May 1910, A. Chevalier 23533 (P04583431, P04583433); Cotonou, 3 April 1962, D. Froment 1198 (BR0000017458450); [Abomey-]Calavi, 6 April 1988, Sinsun 608 (BR0000000054513, BR0000017458443); Bourgou province, Kalale, 10°15'N, 3°23'E, 380 m, 16 October 1988, J. Lejoly 88/194 (BR0000000545171); Davougon, 7°10'N, 1°57'E, 9 October 1996, M. Ayichedehou 69 (BR0000000545898); Bourgou, Malanville, Sota, 11°47'N, 3°23'E, 13 November 1998, V. Adjakidje et al. 2575 (BR0000017458436); Yaoui (Toui-kilibo), 8°28'N, 2°37'E, 23 July 1999, Oumorou & Lejoly 570 (BR0000000546882); Cotonou, 26 July 1999, F. Genco 177 (BR0000000559288); Ouidah (“Whydah”), P.E. Isert s.n. (C10004313).
BOTSWANA: [North-West District] Kwebe Hills, 4 February 1898, Lugard 153 (K); Central distr., Sephope [Sefhophe], 11 February 2006, Farrington et al. 0316769 (K); [North-West District] Moremi Game Reserve, 22 February 2008, Heath 1537 (K).
BURKINA FASO: Soum, Djibo-Barabole, 4 October 2006, Carre et al. 0353348 (K); Ouagadougou/ Sikasso, 25 September 1958, J.G. Adam 15436-4 (P00695360); Axe Djibo-Barabole à 8 km Ouest-Nord, 14°05'55"N; 1°42'34"E, 4 October 2006, L. Sanou et al. BUR 451 (P00544385); Zabre, 23 July 1975, B. Toutain 1188 (P04583418); Tengongo, 8 km S of Ouagadougou, 12°17'N, 1°31'W, 23 June 1983, J. Lejoly 83/251 (BR0000017458337); 22 km N of Banfora, 10°48'N, 4°42'W, 12 June 1983, J. Lejoly 83/071 (BR0000017458344); Bale, 10 km E of Boromo, 11°46'N, 2°51'W, 5 August 2005, van der Maesen et al. 8026 (BR0000017458351).
BURUNDI (selected specimens): Rutana, 1 March 1960, Breithof 19 (BR0000017460835); Bujumbura, 25 January 1967, Lewalle 1525 (BR0000017460866, K, M); Bubanza prov., Randa, 3 December 1977, Reekmans 6658 (BR0000017460941, K); Bururi prov., Mugara, 27 November 1979, Reeekmans 8356 (K); Bubanza prov., 21 March 1980, Reekmans 8749 (
CAMEROON: Adamawa province: near Koti (50 km SE Banyo), 24 June 1967, R. Letouzey 8713 (P04583439); East province: Bordoukou, September 1947, anonym 1264 (P04583550); Far North region: about 5 km W of Maroua, c. 500 m, 2 September 1964, W.J.J.O de Wilde & B.E.E. de Wilde-Duyfjes 2971 (P04583388, BR0000017458535); 24 August 1964, Letouzey 6389 (P04583487); Waza (Sd Maltam), 19 September 1972, A. Gaston 3201 (P04583377); Kaele (55 km SSE Maroua), 65 km SW of Kousseri, Kousseri-Waza road, 10 September 1984, S. Lisowski B-985 (BR0000017458528); North region: Garoua, 7 September 1964, Bounougou 114 (K, P04583486); Mayo Alim (52 km S Gouna), 9 December 1964, J. & A. Raynal 12406 (P04557321, P04583389); Poli, 28 September 1968, H. Jacques-Felix 8385 (P04583513); West region: km 18 Bafoussam-Foumbot road, 1 km E of Nchi R., 5°17'24"N, 10°18'36"E, 1000 m, 15 December 1971, A.J.M. Leeuwenberg 8903 (BR0000017461092, M; P04583397).
CAPE VERDE: San Nicolau island, 1851, Bolle s.n. (K-001134117); Sal, Fogo [Sao Philippe] & Santo Antao Islands, 1875, Loewe 37 (
CENTRAL AFRICAN REPUBLIC: [Nana-Mambéré prefecture] Bouar, 1914, Midbraed 9385 (
CHAD: Batha region: Mateti c. Kouki, 28 August 1968, A. Gaston 1795 (P04583383); Chari-Baguirmi region: Baguirmi, 10-16 August 1903, A. Chevalier 9571 (P04583489); Ennedi-Ouest region: Mortcha, Koalga, 27 September 1935, B. Zolotarevsky, M. Murat & L. Dupont 497 (P04583447); Logone Occidental region: Deli, 14 December 1962, Mosnier 1517 (P04583570); Moundou, 6 December 1964, Audru 1898 (P04557323, P04557324); Doba, 17 August 1964, Audru 824 (P04583393); Kanem region: Bochoumanga, 12 August 1964, A. Gaston 324 (P04583450); Touil, 12 August 1964, A. Gaston 301 (P04583569); Mayo-Kebbi region: Koyom, 7 July 1969, Fotius 1517 (P04583413, P04583425); Moyen-Chari region: Moussafoyo, 1 July 1964, J. Audru 114 (P04583438); c. Moussafoyo, 25 October 1968, A. Gaston 2307 (P04583384); N’Djamena region: Ndjamena, 25 July 1984, S. Lisowski E-1 (BR0000017458511); Ouaddaï region: Abeche, 26 June 1966, A. Gaston 766 (P04583449).
COMOROS: [no precise location] 1884, Humblot 438 (
COTE D’IVOIRE: Agnéby-Tiassa region: N’douci, 20 July 2006, A. Mangara 218 (BR0000000959397); Bounkani region: Bouna National Park, Tehini, 21 August 1964, R.A.A. Oldeman 287 (BR0000017458313, K, P04583391); Kakpin, 12 August 1967, G.J.H. Amshoff 711 (BR0000017458290); Gbêkê region: Baoule-Nord cercle, between Diahbo and Bouake, 9 July 1909, A. Chevalier 22077 (P04583469); Bouake, 370 m, 1 June 1962, A.J.M. Leeuwenberg 4275 (K, P04583386); San-Pédro region: Bereby [Grand-Béréby], November 1964, G.J.H. Amshoff 646 (BR0000017458320, K); Worodougou region: 2 June 1909, A. Chevalier 21823 (P04583488).
DR CONGO (selected specimens): Bandundu province: Lieu, Kikwit, 21 November 1990, Masens 437 (K, BR0000017459587); Bas-Congo province: Moanda, [no date] Bittremieux 287 (BR0000017458955); Madimba, 13 December 1956, Carlier 359 (K; M); Songololo, 19 December 1959, P. Compere 1101 (BR0000017459143); Équateur province: Befale, 23 June 1958, Evrard 4277 (K); Kasai-Oriental province: Gandajika, 11 December 1956, L. Liben 2081 (BR0000017459815); Bakwanga [Mbuji-Mayi], August 1958, L. Liben 3695 (BR0000017459563); Mweka, 17 June 1958, Decleirq 41 (BR0000017459518); Katanga province: Bukama, December 1948, de Witte 4841 (K); Kongolo, 9 November 1974, Thompson s.n. (K); Kinshasa province: Leopoldstad [Kinshasa], 1922, Achten 7 (BR0000017459020); Kongo Central province: Matadi, 20 December 1887, F. Hens 324 (P04583547); North Kivu province: plain S of Lake Edward, 1100 m, May-June 1929, H. Humbert 8193 (P04583509, P04583423); Katanda, September 1937, J. Lebrun 7726 (P04583549; M); Beni, November 1954, de Witte 11486 (K); Ndemo, Albert [Virunga] National Park, October 1956, de Witte 13617 (K); Orientale province: Bili, May 1931, Lebrun 2825 (K, P04583510); Yangambi, October 1936, J. Vouis 2691 (P04583440); Dungu, Garamba National Park, 2 April 1950, Noirfalis 115 (K); Opala, 13 November 1976, S. Lisowski 43281 (BR0000017459969); South Kivu province: Ruzizi, 1950, Germain 5809 (K).
ERITREA: Northern Red Sea region: Assaorta, August 1902, Pappi 2663 (
ETHIOPIA: Afar region: Awash National Park, 12 April 1969, Gilbert 1210 (K); Shewa, Awash station, 8°59'N, 40°10'E, 1 August 1975, P.C.M. Jansen 2490 (BR0000000569222); Benishangui-Gumuz region: between Guba-Mankush & Bambudi, 28 October 2010, I. Friis & al. 13628 (BR0000015246189); Gambela region: Gambela, 17 May 1976, Ash 3419 (K); Gambela, 15 September 1988, Pavlov 330 (
GABON: [without precise locality], 7 August 1966, Halle & Thomas 364 (P04557343).
GHANA: Greater Accra region: Accra, 14 March 1927, Deighton 583 (K); Prampram, November 1951, Morton 6080 (K); Accra, 23 September 1954, J.O. Ankrah 20022 (P04583503); Kpong, 10 August 1958, J. Lebrun 11329 (BR0000017458368); [Accra], Dome, near Achimota, June 1961, F.R. Irvine 5030 (K; BR0000017458375); [Accra], Legon, 6 April 1963, Dokozi 381 (
GUINEA: Kouroussa, July 1900, M. Pobeguin 369 (P04583545); Nzerekore, S of Mt Yonon, 7°58'12"N, 9°03'W, 446 m, 26 April 2011, C. Jongkind et al. 10331 (P06807575, BR0000017458252); Kankan, 1 November 1966, S. Lisowski 61918 (BR0000017458269).
KENYA (selected specimens): Tsavo Natural Park, 11 April 1966, Gillett 17268 (K); Baringo county: Baringo distr., 31 October 1964, Leippert 5275 (K); Rift Valley, 6 km from Marigat to Kabarnet, 0°28'12"N, 35°58'48"E, 31 October 1964, H. Leippert 5275 (
LIBERIA: Gbarnga distr., Suakoko, 20 April 1952, M.L. Blickenstaff 55 (P04583499, BR0000017458276).
MADAGASCAR: Central Madagascar, August 1880, Parker s.n. (K); valle de Manambolo, rive Gauche, environs d’Isomono, Monts Kotriha et Isomonobe, 1933–1934, H. Humbert 12836 (P04582864; P04582866); Ranomafana, 20 February 1964, Peltier 4748 (P04934053); Alaotra-Mangoro region: D’Ambatondrazaka distr., Alaotra Lake, 1918, Esperandieu 13 (P05196704); Ambatondrazaka, October 1938, Cours 1262 (P04582920); Analamanga region: Antananarivo, [no date], anonym s.n. (
MALAWI: Central region: Chitala, 12 February 1959, Robson 1552 (
MALI (selected specimens): Bamako, June1929, Waterlot 1199 (P04583471); Gao region: Gao, 12 September 1927, Hagerup 334 (BR0000017458191, K); Tabankort, 28 August 1931, A. Leclercq 42.647 (P04583461); Menaka, 27 November 1932, A. Leclercq 813 (P04583459, P04583463); Gao, 1 July 1935, de Wailly 4704 (P04583457); Kayes region: 5 km E Macina (c. Nioro), 22 October 1969, G.Boudet 5973 (P04583373); Kidal region: Adrar [Adrar plateau], September 1969, Popov 69/29 (
MAURITANIA (selected specimens): Assaba region: Tahmeire, Sortie Sud de Kiffa sur la route de Kankossa, 16°32'47’’ N, 11°24'9’’ E, 9 September 2008, J.-N. Labat et al. 3957 (P00577618); Brakna region: between Bassi Nguidi and Mal, 10 October 1960, M. Mosnier 7487 (P04583562); Gorgol region: Rinndiao, 1972, Naegele s.n. (K); Maghana, 22 August 1986, Barriere 45 (P05196644); Trarza region: Tamzak, 19 October 1963, J.G. Adam 19389-4 (P00694379, P00695683); Tagant region: Letfatar, 26 October 1910, M. Chudeau s.n. (P04583502); Moudjeru [Moudjeria], 24 June 1911, M. Chudeau s.n. (P04583501); Nbeika, 27 September 1997, G. Heynemann 2940 (BR0000000938991).
MAURITIUS: Morne Brabant [Peninsula], 23 February 1973, Gueho 15575 (K).
MOZAMBIQUE: Manica province: Manika & Sofala, Chimoio, 23 February 1948, Garcia 324 (K); Dombe, Machonga, 23 November 1965, A. Pereira & A. Marques 834 (BR0000017458924); Nampula province: Erati, Namapa, 21 March 1961, Balsinhas & Marrime 304 (
NAMIBIA: Erongo region: 18 January 1934, Dinter 6850 (
NIGER (selected specimens): Agadez region: Ighazer plain nr Agades, 9 September 1965, Popov 35 (
NIGERIA: Adamawa state: Yola, 19 July 1909, Dalziel 51 (K); Anambra state: Onitsha, 3 May 1944, Onochie 7526 (K); Vogel Peak area, Dawo, 520 m, 19 November 1957, Hepper 1408 (K, P04583508, BR0000017458542); Bauchi state: Katagum distr., 9 July 1907, Dalziel 195 (K); River Benue, September 1910, Talbot s.n. (
REPUBLIC OF CONGO: [Bouenza Department] Loudima, 12 May 1979, Bonnin 401 (K); Brazzaville, July 1908, A. Chevalier 4188 (P04583494); Brazzaville, November 1884, M. Thollon 919 (P04583553); Ngatsou, 55 km NE of Brazzaville, 13 July 1985, G. Cusset 1389 (P04583565).
RWANDA: [Eastern province] Kibungu, 10 April 1958, Troupin 6962 (K, BR0000017460828); [Harare province] Bitshumbi, 1933, G. de Witte 1030 (P04583548).
SENEGAL (selected specimens): Dakar: 18 September 1930, J. Trochain s.n. (P04583442); environ Dakar, 7 August 1948, J.G. Adam 1678 (P00695351); Alamadies, 2 October 1955, J.G. Adam 10958 (P00695339); Diourbel region: Bambey, 13 December 1960, J. & A. Raynal 6680 (P04583430); Kaolack region: Kaolak, 1950-1951, R.P. Berhaut 401 (P04583551); Kédougou region: Niokolo-Koba, 12 June 1958, J.G. Adam 14.345 (P04583484); Louga region: Lagbar, 18 September 1956, J.G. Adam 12344 (P00695336); Gallayel, 8 September 1965, M. Mosnier 2232 (P04583392); Saint-Louis region: near Dagana, July 1828, Leprieur s.n. (P04583582); Ross-Betio [Ross Bethio], 25 August 1965, Audru 2281 (P04583396); Dagana, 12 October 1969, Hepper 3642 (K); Fete-Ole, 15 October 1976, A. Cornet 536 (P04583398); NE of Lac de Guiers, 16°20'N, 15°45'W, 22 November 1984, P. Bamps 7596 (BR0000017458153); Tambacounda region: Tambacounda, 1 October 1988, C. van den Berghen 8188 (BR0000017458177); Thies region: Thies, 5 November 1939, Adam 8944 (M); Nianing, 16 August 1954, R.P. Berhaut 5370 (P04583491); Ziguinchor region: Basse-Casamance [National Park], October 1988, C. Vanden Berghen 8188 (BR0000017458177).
SIERRA LEONE: Sierra Leone, 1798, Afzelius (LINN-HS 152.3); Eastern province: Njala, 25 September 1926, Deighton 2119 (K); Northern province: nr Falaba, 31 March 1892, Scott Elliot 5423 (K); Makeni, 31 August 1928, Deighton 1276 (K); Makump, 4 May 1929, Deighton 1413 (K); Rokupr, 20 February 1950, Jordan 382 (K).
SOMALIA: Bay region: Bur Akaba [Buur Hakaba], Gillett Hemming 24893 (K); Buur Heybe, 1985, O’Brien s.n. (K); Durdur, 1 June 1989, Thulin & Mohamed 6759 (K).
SOUTH AFRICAN REPUBLIC: KwaZulu-Natal province: Zululand, 8 December 1971, Pooley 1552 (E; M); Limpopo province: Waterberg distr., 3 March 1925, Galpin 153 (K); Dongola Reserve, 15 March 1948, Codd 3845 (K); Transvaal [Limpopo] prov., Kruger National Park, 4 February 1949, Codd & de Winter 4983 (K); Transvaal, Soutpansberg, 31 January 1962, Schlieben 9279 (K); Malalane, Kamatipoort [Komatipoort], 20 January 1966, Hilliard & Burtt 3613 (E); Messina [Musina], 8 January 1974, Pienaar 283 (K); Mpumalanga province: Komatipoort, December 1917, Moss 607 (
SOUTH SUDAN: Gondokoro, February 1910, Mearns 3058 (
SUDAN: [Kordofan state] Cordofan Mt., 3 October 1839, Kotschy 119 (
TANZANIA (selected specimens): Arusha region: Lake Manyara, 15 June 1965, Greenway & Kanuri 11860 (K); Dodoma region: Bahi, 3000 ft, 23 April 1962, Polhill & Paulo 2132 (K, BR0000017458801); Kilimanjaro region: Lake Chala crater rim, 3°19'12"S, 37°40'48"E, 8 January 2004, A. Hemp 4242 (
TOGO: Kara region: Kpagouda, 10 May 1980, H. Scholz 870 (K); Maritime region: Lome, 1900-1902, Warnecke 6 (
UGANDA (selected specimens): Central region: Singo co., Wattuba, 13 April 1970, Katende 103 (K); Mukono, Kyagwe, 20 March 1992, Rwaburindore 3395 (K); North Buganda, Mukono, Kyagwe, Bule-Buyaga, 0°17'S, 32°42'E, 1200 m, 20 March 1992, P.K. Rwaburindore 3395 (BR0000017458610); Eastern region: Busoga distr., 17 April 1953, Wood 724 (K); Kampala, [no date] Liebenberg 839 (K); Western region: Toro distr., 24 December 1934, Taylor 2655 (
ZAMBIA: Central province: Chitala, 12 February 1959, Robson 1552 (K); Eastern province: near Changwe, between Petauke & Mwape, 650 m, 16 December 1958, N. Robson 959 (
ZIMBABWE: Manicaland province: Sabi valley, Nyanyadzi, 1800 ft, 3 February 1948, H. Wild 2494 (BR0000017458917); Nyanga distr., 29 November 1959, Goodier 686 (K); Mashonaland West province: Urungwe, 29 January 1958, Drummond 5326 (K); Masvingo province: Nuanetsi, 8 May 1958, Drummond 5702 (K); Chibi distr., 29 December 1962, Moll 358 (E); Matabeleland South province: Beitbridge, 22 March 1967, Mavi 280 (K); Insiza, 8 February 1974, Mavi 1522 (K); Midlands province: Queque [Kwekwe], 1 May 1976, Dye 473 (K).