Glinus lotoides L., Sp. Pl. 1: 463 (1753).

Sukhorukov, Alexander P., Sennikov, Alexander, Veranso-Libalah, Marie Claire, Kushunina, Maria, Nilova, Maya V., Heath, Roger, Heath, Alison, Mazei, Yuri & Zaika, Maxim A., 2021, Evolutionary relationships, biogeography and morphological characters of Glinus (Molluginaceae), with special emphasis on the genus composition in Sub-Saharan Africa, PhytoKeys 173, pp. 1-92 : 1

Glinus lotoides L., Sp. Pl. 1: 463 (1753).

Mollugo glinus ≡ Mollugo glinus A.Rich., Tent. Fl. Abyss. 1: 48 (1847), nom. illeg. superfl. Lectotype ( Adamson: 1961: 127): [icon] Alsine lotoides sicula in Boccone (1674: 21, tab. 11, figure B). Superseded neotype ( Jeffrey 1961: 15): [Italy] Sicily, Boccone (OXF, n.v.). Note. The protologue of Glinus lotoides L. is based on two main elements, an illustrated treatment of Alsine lotoides sicula from Sicily ( Boccone 1674: 21, tab. 11, figure B) and another illustrated treatment of Portulaca baetica , Portulaca luteo flore , Portulaca spuria aquatica from Spain ( Barrelier 1714a: 47, 1714b: figure 336). Linnaeus had not seen any specimen of the species prior to the publication of the protologue ( Adamson 1961), in particular the collection of P. Boccone at OXF ( Jarvis 2007). The herbarium collections of J. Barrelier are no longer extant (cf. Morton 1970) and had seemingly never been consulted by any botanist because all Barrelier’s legacy but drawings was destroyed by fire after his death ( Barrelier 1714a). There were two attempts to lectotypify the name Glinus lotoides . Jeffrey (1961) designated a specimen collected by Boccone and kept at OXF; although this specimen is associated with the illustration in Boccone (1674), it was not examined by Linnaeus and therefore is not part of the original material. Jeffrey’s lectotypification is in effect neotypification. Adamson (1961) designated the illustration in Boccone (1674) as lectotype. This lectotypification is correct and supersedes the neotype designated by Jeffrey.

Holosteum hirsutum = Holosteum hirsutum L., Sp. Pl. 1: 88 (1753). Holotype: India. Ex Malabariae [Malabar Coast], Hb. Van Royen 899 (L, n.v., after van Steenis 1966).

Glinus dictamnoides = Glinus dictamnoides Burm.f., Fl. Ind.: 113 (1768). Lectotype (designated here): [icon] Plate 356, figure 6 in Plukenet (1705). Note. The name G. dictamnoides was erroneously attributed by Fenzl to Linnaeus (1771); however, the latter clearly refers to Burman’s "Flora Indica" where G. dictamnoides was described ( Burman 1768). Burman (1768) cited no specimens in the protologue. Both Burman (1768) and Linnaeus (1771) cited Plukenet (1705) who depicted this plant in the Plate 356, figure 6. This image shows a hairy shoot with the rounded leaves in whorls and almost sessile verticillate flowers. The obovate vs. orbicular leaf shape was the main character known to Burman to distinguish G. lotoides and G. dictamnoides in situ ( Burman 1768; see also a drawing of G. lotoides in the table 36, figure 1). The orbicular and greyish-green leaves of G. dictamnoides were similar to those of Dictamnus creticus Garsault (≡ Origanum dictamnus L.), and such plants were found in Madras [now Chennai], India: "dictamni cretici facie, maderaspatana". Based on the protologue of G. dictamnoides , Merrill (1921) synonymized this species name with G. lotoides . In agreement with Merrill’s opinion, we designate the cited illustration as lectotype and retain Burman’s name in the synonymy of G. lotoides . The specimens seen from India usually have rounder leaves with ± scattered pubescence and shorter perianth (usually 5.5 mm long in fruiting) compared to the European populations, which have obovate and usually hirsute leaves and longer perianth reaching 6.5-8.5 mm in length. These characters were presumably the main argument to consider the Indian plants as G. lotoides subsp. hirtus ( Almeida 1998) based on the name Mollugo hirta described from South Africa but erroneously applied to Indian plants ( Clarke 1879). Mollugo hirta represents the plants with different characters (see also notes under G. hirtus ). The African plants corresponding with G. dictamnoides are present in eastern and southern parts of the continent. It should be noted that the density of pubescence is very diverse in African plants, and those growing in a humid climate (e.g., Nigeria, Cameroon) usually have green leaves with scattered hairs. The intermediate forms in leaf shape and pubescence degree were frequently seen in the herbarium collections. In light of our molecular studies showing a mixed position of G. lotoides and G. dictamnoides , and scarce morphological differences between them, we prefer to synonymize G. dictamnoides with G. lotoides .

Tryphera prostrata = Tryphera prostrata Blume, Bijdr. Fl. Ned. Ind. 11: 549 (1826). Note. Described from two localities on the Island of Java, Indonesia: Pamanukan and Cheribon [Cirebon]. The authentic specimens are not traced, but the description of the genus and species ( Blume 1826) indicates the identity of T. prostrata to G. lotoides .

Pharnaceum pentagynum = Pharnaceum pentagynum Roxb., Fl. Indica 2: 103 (1832). Lectotype (designated here): [without location, year and collector] "Hb. Roxb." (K000641797!). Note. Another authentic specimen was found at BR: "Herb. Roxburghii" (BR0000005227204).

Glinus lotoides virens = Glinus lotoides var. virens Fenzl, Ann. Wiener Mus. Naturgesch. 1: 358 (1836).

Mollugo glinus virens ≡ Mollugo glinus var. virens (Fenzl) Oliv., Fl. Trop. Afr. 2: 590 (1871). Lectotype ( Adamson 1961): South Africa. At the junction of Fish and Orange rivers, left side of the Orange River (ca. -28.1, 17.175): ["Garip, bei Verleptpram, am Ufer des Flusses und in der steinigen Niederung, unter 500 Fuss", 17 Sep 1830,] J.F. Drège (K000232021; isolectotypes HAL0117929, HBG516708, K000232020, S05-5102, S05-5103). Note. In the type citation, the locality information is added from Drège (1843: 92) and the collection date is complemented from Gunn and Codd (1981). When establishing G. lotoides var. virens , Fenzl (1836) cited several validly published plant names as synonyms ( G. dictamnoides , Pharnaceum pentagonum , Physa madagascariensis ), and also illustrations and specimens. Since this new name was at the rank of variety and the synonyms were at the rank of species, and Fenzl provided his own description and used his own original material, his variety does not need to be treated as based on any of the synonyms included. Indeed, Adamson (1961) designated a separate type for this varietal name, a specimen collected by Drège in South Africa and cited by Fenzl (1836) in the protologue. This lectotypification is technically correct and should therefore stand. Fenzl (1836) established his variety for the plants that are less villose (or glabrescent at maturity) than the type variety of G. lotoides , with the perianth 5 mm long, and without petaloids. The distribution area of this new variety was circumscribed as East India, Timor, Arabia, Madagascar, and South Africa ( Fenzl 1836). This variety was accepted by the later authors, and its range was widened to include many regions of tropical Africa ( Oliver 1871 sub Mollugo glinus var. virens ; Adamson 1961; Gonçalves 1970, 1978; Figueiredo and Smith 2008; Klaassen and Kwembeya 2013). However, many sheets of similar looking species ( G. hirtus and G. zambesiacus ) were previously identified as G. lotoides var. virens , and this name was therefore used erroneously in many treatments.

Glinus micranthus = Glinus micranthus Boiss., Diagn. Pl. Orient., ser. 1, 10: 11 (1849). Holotype: [Lebanon] Rascheya [Rachaiya], in ruderatis, May-July 1846, E. Boissier s.n. (G00506579!) located at G-BOIS!

Description.

(Fig. 19 ). Annual, stellate pubescent. Stems prostrate or ascending, sometimes reaching 2 m in length, but usually shorter (up to 70 cm), prickles absent. Leaves grey or light green, usually moderately pubescent, sometimes bicolored (light green adaxially and grayish-green abaxially), entire, crisp or denticulate, obovate, broadly obovate or almost roundish; petioles 2-10(15) mm, blades (10)15-60 × 4-20(25) mm, mucronulate. Flowers in clusters of 4-12, distant; sessile or with pedicels up to 15 mm (in fruiting the pedicels, if present, may be up to 20 mm long); white or yellowish, sometimes rusty inside and white abaxially, mid-vein green; flower buds and anthocarp ovoid. Perianth segments (tepals) oblong or ovate, entire or sometimes denticulate, in flowering 5.0-6.5 mm long, in fruiting 7.0-8.5 mm long, with a mucro 0.5-1.0 mm long; white petals (1-3) sometimes present (usually much shorter than tepals). Stamens (5)10-16[∞], obdiplostemonous or alternisepalous, outer stamen series (petaloids) sterile, terminating with 1-3 teeth; anthers 0.7-1.0 mm long. Stigmas (3)5, 0.9-1.5 mm long (sometimes a style is present up to 1.0 mm long). Seeds 0.45-0.6 × 0.40-0.55 mm, dark red or almost black, colliculate and without longitudinal striae; aril hood easily visible, 0.20-0.30 mm long.

Note.

The five-staminate and three-carpellate individuals of G. lotoides are considered here as abnormal forms that seem to be very rare. We assume that such specimens investigated earlier in relation to the floral formulae by Müller (1908) rather belong to G. hirtus .

The plants with green or glabrescent leaves are more common in the wet climate (e.g., West Africa), and the densely pubescent populations are most frequent in drier conditions (Fig. 20 ).

Habitat.

Sandy sites in river valleys, limestone, clayey and rocky places, seasonally inundated areas; at altitudes of up to 2000 m. This species prefers semiarid regions and only limited number of specimens from West Equatorial Africa are present in the collections.

Distribution

(Fig. 21 ). Angola: [Luanda prov.] Barra do Bengo, Jan 1853, F. Welwitsch 2415 (BM); Luanda distr., between 8 and 9°S, Alto das Cruzes, Sep 1857, F. Welwitsch 2412 (BM, K, LE, M, P04576620); [Namibe prov.] Mossamedes, Cabo Negro, Sep 1859, F. Welwitsch 2416 (BM, K); [Namibe prov.] Mossamedes, Cavalheiros, Jul 1860, F. Welwitsch 2413 (BM); [Namibe prov.] Bero and Giraul rivers, Jun 1900, J. Gossweiler 72 (COI00070553); Luanda, 1903, J. Gossweiler 223 (BM, LISC031995, P04576560); [Cunene prov.] nr Cahama, 18 May 1909, H.H.W.Pearson 2545 & 2548 (BM, BOL217405, K); [Namibe prov.] Mossamedes, Rio Croca, 3 Jun 1937, L.W. Carrisso & F. Sousa 242 (BM, LISC031994); [Namibe prov.] Porto Alexandre, 15 Sep 1955, E.J. Mendes 79 (BM, LISC031997); [Namibe prov.] Mossamedes, Camucuio, 14 Oct 1955, E.J. Mendes 410 (LISC031998, M); Cuanza Sul prov., Foz do Rio Cuvo, 4 Mar 1967, J.B. Teixeira et al. 11054 (LISC032003, WAG0104171); [Huila prov.] Chiange, 4 Dec 1970, J.A. de Sousa 86 (K, LISC032011);

Benin: Zou dept. 8 Mar 1970, L. Aké Assi 11109 (G); [Alibori dept.] Malanville, 13 Nov 1998, V. Adjakidjè 2593 (WAG0235875); [Alibori dept.] Malanville, 11°52'N, 3°23'E, alt. 270 m, 26 Apr 1999, P. Houngnon et al. 6507 (BR0000018268560, K, WAG0235862); Atakora dept., Tanguiéta, 20 Jul 2000, B. Sinsin 3667 (WAG0235863);

Botswana: Ngamiland distr., Kwebe hills, 28 Dec 1897, E. Lugard 73 (K); [Bakgatla tribal region] Mochudi, 24°10'S, 26°05'E, 1914, C.C. Harbor 6584 (BM); [Bakgatla tribal region] Mochudi, Jan 1914, F.A. Rogers 6584 (PRE0403681); [Ngamiland distr.] Maun, Dec 1967, F.L. Lambrecht 449A (K); nr Maphaneng, 19°56.75'S, 23°26.2'E, 4 Dec 1977, P.A. Smith 2115 (K);

Burkina-Faso: [Oudalan prov.] Markoye, 15 Aug 1975, Anonymous 45121 (P04576576); [Sanmatenga prov.] NE of Pissila, 18 Jun 1987, J. Lejoly 87/071 (BRLU0026261); [Oudalan prov.] Oursi, 10 Oct 1988, S. Guinko s.n. (WAG0042080); Oudalan prov., Tin Akoff, 7 Oct 1998, S. Kahlheber 1001 (FR-0108631); Oudalan prov., Yomboli, 24 Jul 1999, J. Müller 82 (FR-0014419); Sourou prov., 50 km W of Tougan on road to Kassoun, 13°10'N, 3°25'W, alt. 276 m, 16 Apr 2002, L. Sanou 100 (K);

Burundi: Bujumbura, plaine de la Ruzizi km 25, alt. 850 m, 28 Sep 1966, J. Lewalle 1046 (K); Bujumbura, 800 m, 28 Sep 1971, J. Lewalle 6145 (BR0000018269406, G, WAG0104170); Bubanza prov., Gihungwe, 800 m, 23 Nov 1974, M. Reekmans 3952 (BR0000018269437, K, MSB-129441); Bubanza prov., Gihungwe, alt. 850 m, 2 Feb 1975, M. Reekmans 4296 (K); Bubanza prov., 3 Dec 1977, M. Reekmans 6677 (K, L0717151, LE, P04576512, PE01696732, WAG0104169);

Cameroon: [Adamawa region] Mayo Lidi, 5 Mar 1933, H. Lhote 82 (P04576622); [Far North region] Dargala, 30 km ESE of Maroua, 21 Aug 1964, R. Letouzey 6314 (K, P04576655, WAG0104162); [Far North region] nr Maroua, 400 m, 13 Sep 1964, W.J.J.O. de Wilde et al. 3240 (BR0000018268638, P04576568, WAG0185134); North region, Garoua, 7 Jun 1975, P. Wit 3056 (BR0000018268607, PE01942337, WAG0317794); [Far North region] Logone-Birni, 9 Jan 1975, D. Dulieu 55720 (P04576563);

Chad: Chari-Baguirmi region, Dar Goulla, 28 Mar 1903, A. Chevalier 7872 (K, P04576585); Tibesti region, Oct 1931, A. Chevalier s.n. (P04576654); [Moyen-Chari region] Iro Lake, 12 Jul 1939, S. de Ganay 127 (P04576532); [Batha region] Mare de Dalato, 31 Aug 1960, H. Gillet 2375 (P04576666); S of Fort Lamy [ N’Djamena], 3 Jan 1963, W.J.J.O. de Wilde et al. 5188 (BR0000018268676, P04576511, WAG1103207); [Logone Occidental region] nr Moundou, 26 Mar 1963, B. Descoings 10624 (P04576565); Mayo-Kebbi Ouest region, ne Leré, 30 Mar 1963, B. Descoings 10727 (P04576567); [Tibesti Ouest Region] nr Zouar, 28 Jan 1965, H. Scholz 58 (B101143617); [Far North region] Delta of Chari, Djimtilo, 29 Jan 1968, J. Léonard 4419 & 4420 (BR0000018268614; BR0000018269505, P04576643); [Tandjile region] Satégui, 9 Dec 1968, Anonymous Fotius 1281 (P04576513); N’Djamena, 28 Jul 1984, S. Lisowski B-39 (WAG1103213); N’Djamena, 5 Feb 1985, S. Lisowski 1808 (BR0000018268706);

Djibouti: [Tadjourah Region] Andabba [Lake], 6 Jan 1957, E. Chedeville 1756 (FT0007026, P04576543);

DR Congo: [Katanga prov.] Kampunda, 3200 ft, 16 Oct 1950, A.A. Bullock 3430 (B101143607, K); [Haut-Uele prov.] Garamba, 700-800 m, 19 Nov 1952, Anonymous Troupin 74 (K); [North Kivu prov.] Albert Lake, 5 Mar 1954, D. van der Ben 1185 (BR0000018269314, K); Katanga, Mitwaba, 21 Aug 1956, J. Brynaert 520 (K)

Eritrea: [Anseba prov.] Keren, May 1870, O. Beccari 45 (FI059438, FT0007013); Dahalak [island] Mar 1892, A. Terracciano 819 (FT0007080); [Southern Red Sea Region] Adarte, 26 Jan 1893, A. Terracciano & A. Pappi 2741 (FT0007022); [Gash-Barka region] Barea Agordat, 27 Jan 1893, A. Terracciano 2852 (RO); [Anseba prov.] Bogos, 3 Feb 1893, A. Terracciano & A. Pappi 2598 (FT0007023); [Gash Barka Region] Mansura, 9 Sep 1905, A. Pappi 6400 (FT0007016); [Gash-Barka region] Damtai, 30 Jan 1909, A. Pappi 7840 (FT0007003); [Gash Barka Region] Mansura, 10 Feb 1909, A. Pappi s.n. (LE); [Gash-Barka region] Barea Agordat, 23 Feb 1909, A. Fiori 978 (FT0007029); [Gash-Barka region] Scetel, Dec 1905, A. Pappi 6747 (RO); [without exact location] 1906, A. Pappi 6834 (RO); [Gash-Barka region] Mansura [subregion], Beni-Amer, 10 Feb 1909, A. Pappi s.n. (LE); Barca [Baraka] river, 30 Mar 1917, I. Baldrati 230 (FT0007036); Keren, 1300 m, 20 Feb 1919, A. Fiori 975 (FI059443; FT0007025); [Gash Barka Region] Setit, 4 Mar 1923, Exped. Corni-Calciati-Bracciani s.n. (FT0007014); [Northern Red Sea region] Wekiro, 50 ft, 31 Jan 1960, D.J. Greathead 139 (BM);

Ethiopia: [Amhara region] Magdala [Amba Mariam], 27 Apr 1862, Steudner 524 (LE); Ogaden [Region] 1891, L. Robecchi Bricchetti 400 (FT0007010); Addis Ababa, 3 Mar 1916, L. Buscalioni 1415 (FT0007002); [Oromia region, Borena zone] Melka Guba, 1937, Anonymous Cutodontis 130 (W); [Southern Nations, Nationalities, and Peoples’ Region] Lago Margherita [Lake Abaya] 21 Jan 1938, A. Vàtova 1582 (FT0007000); [Southern Nations, Nationalities, and Peoples’ Region] Gondaraba, 29 May 1939, R. Corradi 7107 (FT007088); [Southern Nations, Nationalities, and Peoples’ Region] Asile, 1 Jul 1939, R. Corradi 7109A (FT0007087); [Southern Nations, Nationalities, and Peoples’ Region] Murle Lake, 18 Jul 1939, R. Corradi 7113 (FT0007085); Somali region, Kelafo, 24 Aug 1960, Anonymous s.n. (K); Addis Ababa to Debre Markos, Blue Nile, 9 Feb 1966, E.F. Gilbert 51 (BR0000018268744, K); [Oromia region, West Shewa zone] Bako, alt. 1650 m, 21 Oct 1969, C. Parker 198 (K); [Gambela region, Anuak zone] Gambela, 1768 m, 6 Jan 1970, J.W. Ash 167 (K); Sidamo prov., 1550 m, 2 Mar 1971, J.J.F.E. de Wilde & M.Anonymous Gilbert 401 (WAG0102276); [Oromia region / Southern Nations, Nationalities, and Peoples’ Region] Ziway Lake, Feb 1973, O. Polunin 11584 (K); [Oromia region, Shewa zone] Meki, 6 May 1976, M.Anonymous Gilbert & T.Anonymous Jefford 4252 (K); [Oromia region, East Welega zone] between Nekemte and Gimbi, 18 May 1976, P.C.M. Jansen 6355 (BR0000018268737, FR0104475, FT0007032, G00350481; MBM369175 - image seen! WAG0181906); [Oromia region, Borena zone] 45 km W of Yavello towards Teltele and Konso, 975 m, 15 Nov 2010, I. Friis et al. 13681 (K, WAG1950295); [Oromia region / Ethiopian Highlands region] Shala Lake, 20 Feb 1979, M.Anonymous Gilbert & S.B. Gilbert 1173 (K); [Gambela region] N of Abobo, Alwero river, 650 m, 20 Apr 1982, I. Friis et al. 2476 (K); [Oromia region, Borena zone] 5 km N of Yabelo, 1600 m, 11 Dec 1998, I. Friis et al. 9418 (K, WAG0322893); [Afar region] 10 km E of Asaita, 400 m, 17 Sep 2001, I. Friis et al. 10345 (K, WAG0394466);

Ghana: [Upper East region] nr Bawku, 12 Dec 1950, C.D. Adams & AnonymousK. Akpabla 4290 (K, P04576600, WAG0185140); [Northern region] 53 miles N of Tamale, 20 May 1952, J.K. Morton 7200 (K, WAG0185141); [Savannah Region] Yapei, 30 Mar 1956, C.D. Adams 3917 (K); Yeji, 11 Apr 1964, J.B. Hall 1251 (K);

Guinea: [Kouroussa pref.] Kouroussa, Dec 1900, M. Pobeguin 605 (K, P04576530); [ Nzérékoré Region] Ouamadou, Dec 1917, J. Berhaut 930 (BR0000018268423); Kankan region, Kankan vill., 23 Apr 1967, S. Lisowski 61907 (BR0000018268508);

Kenya: Makueni county, Kibwezi, 975 m, without date, P. Luke 14330 (K); [Turkana county] Lokitaung, 3000 ft, Jan 1932, A.M. Champion 14 (K); [Eastern prov.] Athi plains, 21 Aug 1938, P.R.O. Bally 7442 (K); South province, Magadi Road, 3000 ft, 8 Aug 1943, P.R.O. Bally 2679 (G, K); [Eastern prov.] Athi River Station, 24 Aug 1947, A. Bogdan 1109 (K); [Marsabit county] 16 miles from Laisamis on road to Marsabit, 2500 ft, 3 Oct 1947, P.R.O. Bally 5471 (K); [Kajiado county] 1 mile S of Kajiado, 1650 m, 21 Feb 1953, R.B. Drummond & J.H. Hemsley 1249 (B101143601, BR0000018268799, FT0007096, K); [Lamu county] Witu, [without date] Thomas 127 (BR0000018269000, G); [Taita-Taveta County] Voi, Sala Gate Rd, 1100 ft, 2 Jan 1967, J.Anonymous Greenway 12937a (K); [Turkana county] Ayangyangi swamp, 1900 ft, 4 Sep 1968, O.M. Mwangangi & D. Gwynne 1235 (BR0000018268829, K); Turkana county, between Lothagam and Kerio delta, 1200 ft, 29 Aug 1968, O.M. Mwangangi & D. Gwynne 1218 (K); [Turkana county] Ayangyangi swamp, 1°55'N, 36°5'E, 12 Jun 1970, B. Mathew & M.D. Gwynne 6768 (FT0007099, K); North Eastern prov., 57 km WS of Wajir, 225 m, 6 Mar 1974, Bally & Carter 16590 (K); Mughwango South swamp, 15 Apr 1972, J. Ament & F.C. Magogo 9 (K); K4, Tana River county, Kora National Reserve, 18 miles from Kora Res. Camp to Asako, 28 Aug 1983, J.Anonymous Mutangah 139 (K); Tana River county, Baomo vill., 30 m, 12 Mar 1990, Q. Luke et al. 141 (K); Embu county, Riakanau, 1125 m, 29 Nov 2000, S.A.L. Smith et al. 253 (K); Isiolo county, Ngare Mara, 0°30'N, 37°39'E, 945 m, 12 Aug 2005, C. Obunyali et al. 261 (K); Machakos county, Kitanga, 1°31'S, 37°11'E, 1660 m, 26 Jan 2005, J.M. Muasya 482 (K); Marsabit county, Log-Logo, 1°59'S, 37°54'E, 520 m, 9 Feb 2005, J.M. Muasya et al. 505 (K); [Coast prov.] Tana delta, 2°24'S, 40°18'E, 5 m, 4 May 2011, C. Leauthaud et al. 160 (K);

Malawi: Nsessi, Dec 1887, L. Scott s.n. (K); Malawi Lake, 1893-1894, AnonymousF. Scott Elliot 8403 (K); [Southern region] Mulanje, 16 Nov 1955, Anonymous Jackson 1761 (BR0000018269536, K); Central region] Ncheu, 29 Sep 1970, A.J. Salubeni 1491 (K); Central Region, Ntchisi Distr., 1060 m, 20 Jun 1970, R.K. Brummitt 11583 (K); Northern region, Karonga Distr., Kaporo, 1550 ft, 2 Jan 1974, J. Pawek 7726 (K); [Southern region] Mangochi Distr., Lake Malawi, 22 Dec 1984, I.H. Patel et al. 1736 (K); [Northern region] Nkhata Bay Distr., S of Mbuzi Hill, 24 Aug 1984, A.J. Salubeni & W.Nachamba 3875 (K); [Central region] Salima Distr., Lifidzi breeding centre, 17 Nov 1985, I.H. Patel & R.B. Kwatha 2898 (K); Southern region, Phalombe, 15°46'47"S, 35°48'44"E, 721 m, 15 Apr 2008, H.T. Chapama et al. 830 (K);

Mali: [Sikasso region] Zamblara, 16 Jun 1899, A. Chevalier 991 (P04576672); [Niger Delta region] Djenné, 30 Jun 1899, A. Chevalier 1129 (P04576675); [Mopti region] Mopti, Niger river, 15 May 1927, O. Hagerup 72 (K); [Timbuktu region] Timbuktu, Jun 1927, O. Hagerup 123a (LE); [ Ségou Region] Macina [The Inner Niger Delta], 14 May 1932, O.B. Lean 14 (K); [Mopti region] Dallah, 19 Jun 1932, J. Rogeon 388 (P04576669); [Mopti region] Sangha, 14 Jan 1938, S. de Ganay 73 (P04576552); [Mopti region] Saré Dina, 25 Mar 1952, J.T. Davey 62 (K); [Mopti Region] Dogo, 7 May 1952, J.T. Davey 102 (K); [ Ségou Region] Kara, 15 Jul 1954, J. T. Davey 192 (K); [Timbuktu region] nr Timbuktu, 6 Aug 1959, Anonymous Popov 90 (BM); [nr Bamako] Korofina, 26 Feb 1960, J. Raynal & A. Raynal 5514 (P04576662); [Sikasso region] Sanzana, 3 Apr 1970, N. Diarra 590 (P04576520); [Timbuktu region] Ngoro, nr Niafunké, 20 Jan 1991, A. Raynal-Roques 23030 (P04576559); [Koulikoro Region] Koulikoro, Niger river, 6 May 1993, R. Ehrlich 463 (B100048300); Kayes region, Comitradougou, 100 km from Diéma, 14°25'N, 8°18'W, alt. 317 m, 25 Feb 2006, R. Dembele et al. ML-283 (K001394833);

Mauritania: Tagant region, May 1937, A. Chevalier s.n. (P04576594); Nouakchott, 13 Feb 1957, J.Anonymous Adam 12980 (P00695348); [Gorgol region] between Kaédi and Mbout, 14 Jun 1960, M. Mosnier 590 (P04576667); [Gorgol region] Rinndiao, 16 Oct 1972, A. Naegele 86 (K); [Trarza region] R’Kiz Lake, 15 Sep 1975, D. Dupont s.n. (BR0000018268485); Tagant region, 32 km SE of Tidjikja towards El Khedia, 25 Oct 1987, M. Carriere et al. 85209 (K);

Mozambique: Gaza prov., 25 km from Vile de João Belo [Xai Xai], 7 Oct 1925, Anonymous Pedro 218 (K); Sofala prov., Gorongosa NP, 12 Oct 1944, F.A. Mendonça 2458 (K); Tete prov., Mazowe River, 1000 ft, 22 Sep 1948, H. Wild 2598 (K); Tete prov., Ilha Micune, 19 Jun 1949, L.A. Grandvaux Barbosa & F. de Lemos 3187 (K); Gaza prov., Guija, between Aldeia do Guija and Aldeia da Barragem, 15 Nov 1957, L.A. Grandvaux Barbosa & F. de Lemos 8141 (K); [Gaza prov.] between Chibuto and Canicado, 11 Oct 1957, L.A. Grandvaux Barbosa & F. de Lemos 8001 (K); [Faza prov.] Sul do Save, Chibuto-Baixo Changana, 24 Aug 1963, A. Macedo & L. Macuacua 1139 (K); Lorenço Marques [Maputo] prov., Namaacha, 5 Jul 1967, A. Marques 2043 (WAG1103167); Gaza prov., Caniçado, 23 Aug 1969, M.F. Correia & A. Marques 1181 (E, WAG1103169); [Sofala prov.] Gorongosa NP, Jul 1970, K.L. Tinley 1938 & 1941 (BR0000018269178, M, WAG1103228); Gaza prov., Limpopo river, 2 Aug 1973, M.F. Correia & A. Marques 3156 (WAG1103171); [Tete prov.] Cahora Bassa, 2 Nov 1973, M.F. Correia & al. 3700 (WAG1103172); [Sofala prov.] Gorongoza NP, 9 Nov 2007, P. Ballings & B. Wursten 945 (BR581201); Cabo Delgado prov., Ruvuma, 15 Nov 2009, Q. Luke 13804 (K, P04557299);

Namibia: Waterberg region, Quickborn, 28 Oct 1928, R.D. Bradfield 301 (K); [Kunene Region] Omuramba to Otamako, 27 Feb 1939, O. Volk 1201 (M); [Khomas Region] Windhoek, Adodamm, Jan 1947, Wiss 826 (M); [Erongo region] Brandberg, Ugab river, 8 Aug 1948, R.Anonymous Stray 2407 (K, PRE0403677); [Erongo region] Kuiseb river, nr Homeb, 11 Oct 1961, W. Giess 3762 (M); [Kunene region] Kaokoveld, Kunene river, 17°15'S, 12°16"E, 16 Aug 1956, R. Story 5826 (K, PRE0403678); [Karas Region] Sendlingsdrif, Orange river, Aug 1963, H. Merxmüller & W. Giess 3259 (M); [Otjosondjupa region] Otjosondu, Omusema river, 1300 m, 10 Mar 1967, R. Seydel 4495 (B101143620, K, M, PRE0614464); [Karas Region] Lüderitz-South distr., Sendlingsdrif, 21 Sep 1972, H. Merxmüller & W. Giess 28689 (M, PRE0403676, WAG1103212); [Khomas Region] Windhoek, Farm Regenstein, Dec 1979, W. Giess 15418 (M);

Niger: [Zinder region] Tasker, [without date] P. de Fabregues 8834 (P04557298); [Tahoua region] Madaoua, Feb 1932, A. Chevalier 43716 (PP04576596); [Tilaberi Region] Dolbel, 24 Oct 1962, J. Jangoux 1214 (BRLU0026263); [Agadez Region] 19°18'N, 7°15'E, 4 Dec 1965, C.F. Hemming s.n. (K); [Agadez region] Touaret, 20°20'N, 07°00'E, 10 Dec 1965, Anonymous Popov 138 (BM); [Diffa region] Bosso, 20 Jun 1966, J.P. Lebrun 13836 (P04576587); [Agadez region] Iferouane, 19°10'N, 08°30'E, 3000 ft, 22 Jan 1970, R.A.H. Davies 32 (BM); [Tahoua region] Abalak, 12 Jan 1975, P. Lavie 870 (P04576521); [Agadez region] Iferouane, 17 Mar 1979, Dulieu 49673 (P04576588); [Agadez region] Azanyares, 18°9'N, 9°17'E, 19 Mar 1979, J.E. Newdy ZP64 (K); [Agadez region] Tarouadji, 30 Mar 1980, E. Schulz s.n. (B101143619); [Agadez region] 100 km S of Agadez, 23 Nov 1985, C. Pase 3149 (K); Niamey, 11 Apr 1987, N. Leman 83 (BR0000018268515); [Tilaberi region] Tapoa zone, West Niger NP, 7 Apr 1996, H. Breyne 6186 (BR0000018268522, WAG0330951);

Nigeria: [Benue State] Abinsi, 1912, J.M. Dalziel s.n. (BM, BR0000018268539, P04576604); [Kwara State] Jebba, 9 Dec 1927, O. Hagerup 699 (K); [Bauchi State] Bauchi, Feb 1929, H.V. Lely 144 (K, P04576593); [Sokoto State], Sokoto, 20 Apr 1946, R.W.J. Keay 15877 (K); [Yobe State] Gashua,8 miles E of Zurmi, bed of Fafara river, 21 Jun 1947, C.F. Onochie 23365 (K); [Niger State] 10 miles N of Bussa, Shagunu, 10°20'N, 4°25'E, 25 Jul 1965, C.D.K. Cook 408 (K); [Niger State] Bida distr., at the bank of Niger river, 1 Mar 1968, B.O. Daramola & A. Binuyo 61946 (K); [Zamfara State] Kaura Namoda area, 22 Jul 1969, M.Anonymous Latilo 62605 (K); [Taraba State] Sardauna, nr Gashaka, 25 Mar 1970, J.B. Hall 1577 (K); [Taraba State] Gashaka distr., Gashaka river, alt. 1100 ft, 25 Mar 1970, Z.O. Gbile & B.O. Daramola 23888 (K); [Niger State] Kainji Lake, 21 Jul 1973, R. Linnavuori s.n. (H1209977); [Kwara State] nr Jebba, 15 May 1974, Latilo et al. 69359 (K, WAG0185138); Borno State, Maiduguri, E. Denys 740 (BR0000018268553, WAG0104167); Borno State, Gajiganna, 23 Feb 2004, S. Kahlheber 1082 (FR-0034796);

Senegal: [without exact location] May 1825, Roger 66 (K); [without exact location and date] Perrottet s.n. (RO); [without exact location and date] herb. Maire s.n. (RO); Galam, [without date and collector] s.n. (P04577294); [Saint Louis region] Richard Toll [village], 26 Feb 1965, Anonymous Roberty 16845 (G); Djibelor, 17 May 1982, C. Vanden Berghen 4987 (BR0000018268478);

Somalia: Somaliland, 75 km S of Hargeisa, near Dofar, 19 Oct 1954, P.R.O. Bally 10115 (G); [Lower Juba prov.] Jubbada Hoose, Badana Lake, ca. 75 km WSW di Chisimaio, 0°32'S, 41°58'E, 28 Oct 1971, Anonymous Moggi & R.Bavazzano s.n. (K); [Shabeellaha Dhexe prov.] Mahaddey Uen [Mahadday Weyne], 13 Mar 1975, R. Bavazzano 442 (FT0007076); [Lower Shebelle prov.] Afgooye-Merca road, 18 Mar 1975, R. Bavazzano 486 (FT0007077); Middle Juba prov., Buale to Baardheer, 29 Jun 1978, S.M.A. Kazmui & al. 777 (WAG0338795); [Middle Shabelle region] 35 km N from Jowhar, 14 Oct 1983, P. Kuchar 15319 (K); [Middle Shabelle Region] Jowhar, 2007, A. Sheik A2007 (K);

South Africa: [Western Cape prov.] Cape of Good Hope, [without date] Drège s.n. (P04576613); [Limpopo prov.] Messina [Musina], [without date] F.A. Rogers 19446 (G, Z-000092197); River Limpopo, nr Messina [Musina], 27 May 1927, R.AnonymousH. Young 14664 (BM); [Zululand] Ubombo, Mkuzi Game Reserve, 100 m, 9 Dec 1959, C.J. Ward 3352 (K, M, PRE0403692, W); [KwaZulu-Natal prov.] Ingwavuma Distr., 12 May 1965, Vahrmeyer & Tölken 977 (K, PRE0403694, STU); [KwaZulu-Natal prov.] Ingwavuma, 11 Jul 1974, H. Furness 97 (E); [Limpopo prov.] Soutpansberg, Limpopo river, 4 Apr 1983, C. Straub 173 (PRE0653966); Limpopo prov., nr Kruger NP, 350 m, 24 Nov 2000, N.H.Anonymous Jacobsen 2431 (BR945920);

South Sudan: [Upper Nile State] bank of the Khor Geyni, Pibor river, 4 Jun 1929, N.D. Simpson 7032 (K); Jonglei State, 8 km S of Maar, alt. 450 m, 18 Feb 1980, J.M. Lock 80/26 (K);

Sudan: Khartoum, 1860, Hartmann s.n. (LE); [River Nile State] Shendi, 25 Oct 1868, K. Schweinfurth 736 (K); “Kordofan” [former province], Dec 1878, Exp. Prout 467 (P04577103); Darfur [Region], Jan 1879, Anonymous 84 (K); [White Nile State] Ed Dueim, 10 Jan 1906, P.L. de Vilmorin s.n. (BR0000018268720); Khartoum, 11 Jan 1933, Anonymous Aylmer 275 (K); Kassala State, Tokar delta, 17-23 Nov 1936, F.W. Andrews A220 (K); [Gadaref State, south of Hawata town], Huweishrat el Qardud forest, 16 Apr 1961, J.K. Jackson 4143 (K); Northern State, Wadi Halfa, 29 Sep 1962, A. Pettet 71 (BR0000018268652, K); Kassala State, Ashkeit, 9 Oct 1962, T. Ahti 16731 (K); Khartoum, Buni, 30 Jan 1963, A. Pettet 107 (K, P04577283); [West Darfur State] Jebel Marra, Zalingei, alt. 3400 ft, 7 Feb 1964, AnonymousE. Wickens 1193 (K); [Central Darfur State] Zalingei, Wadi Aribo, 18 Jan 1965, W.J.J.O. de Wilde et al. 5348 (WAG1103211); Khartoum, 12 Mar 1965, W.J.J.O. de Wilde et al. 5836 (BR0000018268713, WAG1103209); North Darfur, Wadi Magrur, 8 Dec 1991, N. Altmann 347 (B100597466);

Tanzania: Kilimanjaro, 1000 ft, Jan 1894, Anonymous Volkens 1642 (G); [Rukwa region] Rukwa Lake, [years] 1898-1900, W. Goetze 1109 (BR0000018268836, G, L1693136, P04577300); [Morogoro region] Kilosa, 28 Dec 1921, C.F.M. Swynnerton s.n. (BM); [Dodoma Region] Kondoa distr., 8 Jan 1928, B.D. Burtt 1869 (K); [Tanga / Manyara regions] Mgera to Kibaya, 1500 m, 23 Aug 1932, Geilinger 1589 (K); [Dodoma region] Mpwapwa to Gulwe, H.E. Hornby 390 (K); [Iringa Region] Ilula, 24 Oct 1932, Geilinger 3292 (K); [Lindi region] 40 km W of Lindi, Lutamba Lake, 3 Oct 1934, H.J. Schlieben 5427 (BM, BR0000018268959, G, M, P04576499); [Shinyanga / Tabora regions] Shinyanga-Nzega distr., Manyonga river, 3500 ft, Oct 1935, B.D. Burtt 5232 (BM, BR0000018268997, K); [Iringa region] Idodi, Oct 1936, E. Ward 29 (K); [Morogoro region] Morogoro distr., Nov 1949, S.R. Semsei 2868 (BR0000018268867, BR0000018268874, K); [Kilimanjaro region] Mwanga, 18 Aug 1951, R.E.S. Tanner 393 (K); [Dodoma Region] Mpwapwa, 3000-3500 ft, 3 Nov 1951, H.E. Hornby 390 (K); [Arusha Region] between Arusha and Moshi, 3500 ft, 23 Sep 1952, P.R.O. Bally 8325 (K); [Tanga Region] Lushoto distr., 1 May 1953, R.B. Drummond & J.H. Hemsley 2346 (B101143608, FT0007097, K); Rukwa Region, Rukwa, 2800 ft, 4 Dec 1954, H.M. Richards 3516 (BR0000018268928, K); [Pwani region] Ruifiji distr., Utete, 2 Dec 1955, E. Milne-Redhead & P. Taylor 7460 (B101143606, BR0000018269024, K); [Katavi region] Mpanda distr., Kibwezi, 3000 ft, 20 Aug 1958, J. Newbould & T. Jefford 1687 (K); [Rukwa region] Rukwa, 21 Oct 1959, H.M. Richards 11517 (K); Dodoma region] Mtera, 19 Apr 1962, Polhill & Paulo 2077 (B101143604, BR0000018268904, K); [Manyara region] Mbulu distr., Lake Manyara NP, 5 Mar 1964, P.J. Greenway & Kanuri 11306 (K); Tanga Region, Korogwe distr., 17 Sep 1965, Semsei 3985 (K); [Manyara region] Tarangire NP, 914 m, 29 Nov 1969, M. Richards 24827 (K); [Dodoma region] nr Dodoma, 3000 ft, 16 Dec 1969, W.J. Mapunda & M.D. Raya 1051 (K); [Arusha region] Engaruka valley, 3530 ft, Oct 1970, H.M. Richards & S. Arasululu 26638 (BR0000018268942, K); [Iringa region] Ruaha NP, 2700 ft, 21 Oct 1970, H.M. Richards & Arasululu 26297 (K); Arusha Region, Masai distr., Magadini vill., 700 m, 8 Aug 1974, B. Mhoro & I. Backeus 2028 (K); [Arusha region] Ngorongoro Conservation area, 1 Oct 1977, J. Raynal 19326 (BR0000018268881, K, P04576664); Morogoro region, Mikumi NP, 500 m, 16 Oct 1986, A. Borhidi et al. 86150 (K); [Kigoma region] Kigoma distr., 22 Aug 1988, anonymous 1687 (K); [Kilimanjaro region] between Moshi and Himo, 920 m, 3 Feb 1996, A. Hemp 945 (P000869013);

Note. Plants of both specimens of Glinus lotoides and G. oppositifolius are present on the same herbarium sheet (Tanzania, Polhill & Paulo 2077, K).

Uganda: [Nakapiripirit distr.] Kakamongole, at base of Mt Debasien, Jan 1936, W.J. Eggeling 2607 (BR0000018268768, K); Karamoja [sub-region], Kidepo river, 3400 ft, Apr 1960, J. Wilson 986 (BR0000018268775, K);

Zambia: Lunsemfwa river, 24 Apr 1929, J. Burtt Davy 897 (K); [Southern prov.] Makambola Farm, Mazabuka, 3300 ft, 28 Apr 1931, C.F. Trapnell 345 (K); [Eastern prov.] Nsefu Game Camp, 750 m, 15 Oct 1958, N.K.B. Robson 138 (K); [Eastern region] Luangwa valley, 13°S, 32°E, 2000 ft, 29 Nov 1957, Stewart 70 (K); [Eastern prov.] Luangwa Game Reserve, Chilongozi Pontoon, 600 m, 11 Oct 1960, H.M. Richards 13333 (K); Zambesi river, Feira [Luangwa], 27 Sep 1962, A. Angus 3350 (K); [Southern prov.] Mazabuka, 2 Oct 1963, H.J. van Rensburg 2515 (K);

Zimbabwe: [Mashonaland] East prov., Mutoko, Mkota Reserve, 1600 ft, 1 Oct 1948, H. Wild 2676 (BR0000018269147, K); [Masvingo prov.] Nuanetsi, nr Malipati, 2 May 1961, R.B. Drummond & R.O.B. Rutherford-Smith 7676 (K); [Matabeleland North prov.] Siabuwa, 2500 ft, Oct 1955, R.M.Davies 1385 (K); [Mashonaland East prov.] Mutoko Distr., Nyangombe river, 1600 ft, 1 Oct 1948, H. Wild 2676 (K); [Masvingo prov.] Chiredzi, 31 May 1971, J.F. Ngoni 147 (K); [Mashonaland West prov.] Lake Kariba, Kessessee Bay, 483 m, 5 Aug 1983, P. Denny 1215 (L0717097);

General distribution.

Africa (Cape Verde, North African countries, Seychelles, Madagascar, Mauritius); W, S & SE Asia; Australia. The distribution in North America is not confirmed in our study.

Figures

Figure 19. Glinus lotoides A an overview of the plant B close-up of the leaves and buds C close-up of the flower D closed anthocarp. Photographer - Ridha El Mokni (Sidi Mechrig - Sejnane, Bizerta Governorate, Tunisia, 23 Oct 2015).

Figure 20. A herbarium specimen of Glinus lotoides (Kayes region, Comitradougou, 100 km from Diema, 14 ° 25 ' N, 8 ° 18 ' W, alt. 317 m, 25 Feb 2006, R. Dembele et al. ML- 283, K 001394833). Copyright of the Board of Trustees of the Royal Botanic Gardens, Kew.

Figure 21. Distribution map of Glinus lotoides in Sub-Saharan Africa (colored in yellow).