Annual, rarely perennial, prostrate herbs or erect subshrubs with a rootstock, covered with stellate or simple (multiseriate, soft and crispate) hairs, in the latter case additionally with multiseriate, stout thick-walled and broad-based hairs (prickles). Stems branched from the base, often forming mats, rarely erect (G. orygioides). Leaves in false whorls, lanceolate to obovate, entire or denticulate (mostly in their upper half), with several lateral nerves that can be adaxially recessed and abaxially prominent. Flowers usually several to many (up to 20) in leaf axils forming loose or rarely dense inflorescences, bracteate, sessile or pedicellate. Perianth of 5 free oblong, ovate or roundish segments, green (brown) abaxially and white, pinkish or yellowish adaxially, with a green or brown midvein, horizontally spreading when fully opened, sometimes white petaloid staminodes present, always shorter than perianth segments. Stamens (2–4)5–30, outer stamen series corresponding to another staminode whorl (if stamen number is more than 5) often with filaments terminating with teeth (and without anthers), 0.3–1.8 mm, oblong or roundish; anthers yellow; pollen tricolpate (studied in G. lotoides: Perveen and Qaiser 2000). Stylodia 3–5, free or united in lower half into a style. Anthocarp (fruit enclosed by perianth) cylindrical or ovate to roundish. Fruit a hydrochastic loculicidal capsule. Ovary three- or five-carpellate, ovules arranged in two rows. Seeds usually more than 50, yellow, red, brown or black, up to 1.0 mm long, ovoid or reniform, smooth or with numerous colliculae; seed aril divided into two parts: a white, very noticeable hood covering the funiculus and a large ribbon-like appendage, sometimes the hood is reduced. Embryo curved; perisperm abundant and easily visible (in larger seeds) or scanty (in small seeds).

The basic chromosome number in G. radiatus is x = 9 (Lane and Keil 1976), which corresponds with that of other Molluginaceae (Bogle 1970, with references therein). However, Mitra and Datta in Löve (1967) indicated the basic number x = 18 for the Indian populations of Glinus lotoides and G. oppositifolius.

(1) The so-called “Norton Shaw valley” lies close to Ngami Lake (Passarge 1904), and “Koobie” should be applied to “Chobe”; (2) The type specimen indicated by Adamson (1961) as kept at GRA is absent in this herbarium (Tony Dold, pers. comm.).

(Fig. 13A–G). Annual or perennial, prostrate or ascending, up to 12 cm tall, often forming large mats up to 1.5 m in diameter. Stems covered with simple crispate hairs; stems, peduncles and sometimes leaf petioles and midveins additionally covered with small, stout hairs (prickles). Leaves rosulate, short-lived, and cauline, green, stout, entire, 10.0–17.0 × 2.0–4.0 mm, oblanceolate or narrowly oblong, sessile or short-petiolate (petioles up to 3.0 mm), mucronulate, veins neither recessed adaxially nor prominent abaxially. Flowers in clusters of 2–7, distant, pedicellate, pedicels 5.0–16.0 mm, in fruiting up to 20.0 mm; buds and closed anthocarp cylindrical. Perianth segments in flowering 6.0–8.0 mm × 2.5–3.0 mm (10–16 mm in diam.), in fruiting 7.0–9.0 mm long, glabrous or sparsely pubescent, dorsally green or with white margins, ventrally white, creamy, pink or pale mauve. Stamens 10–15, outer stamen series (petaloids) sterile, with filaments terminating in teeth; anthers 1.3–1.5 mm long. Stigmas 3, ~1.2 mm long. Seeds 0.6 × 0.5 mm, almost black, with longitudinal ridges carrying colliculate cells; aril hood clearly visible, 0.25 mm long.

In the herbaria G. bainesii (Fig. 14) is often confused with G. oppositifolius, but it is differentiated from the latter species by having small prickles, larger perianth and anthers, and seeds with longitudinal ridges (Table 5).

Riverbed sands, flat flood plains, margins of muddy seasonal pans, sparsely vegetated areas at elevations 0–1200 m a.s.l. Associated plants found in Botswana: Dicerocaryum eriocarpum, Pterococcus oppositifolius var. oppositifolius, poor soil Cypreaceae and Poaceae spp. (A. Heath & R. Heath 734, K); Fuirena pubescens, Neptunia oleracea, Hibiscus meeusei, Cyperus compressus, Senna obtusifolia (A. Heath & R. Heath 428, K); Vahlia capensis, Heliotropium ovalifolium, Cynodon dactylon, Streptoglossa decurrens, Jamesbrittenia elegantissima (A. Heath & R. Heath 1417, K). Flowers during both the early and the main rains.

(Fig. 15). Botswana: Ngamiland, Kwebe Hills, 3300 ft, 4 Feb 1898, E.J. Lugard 152 (K); Northern Bechuanaland, Sigere pan, 30 miles W of mouth of Nata river, 896 m, 25 Apr 1957, Drummond & Seagrief 5216 (K); Ngamiland, Sehitwa, Lake Ngami, 930 m, 25 Mar 1961, H.M. Richards 14849 (K); Northern distr., 17 km SE of Maun at Samadupe drift over Botletle river, 23 Jan 1972, H. Biegel & G. Russell 3733 (BR0000017454490, K001394831, M, P04577251); Northern distr., Moremi reserve, 19°10.4'S, 23°15.7'E, 26 Jan 1974, P.A. Smith 848 (B101143636, K, WAG1103318); Northern distr., Nata river delta, 21 Apr 1974, J.F. Ngoni 529 (K, PRE0825248); Ngami Lake, 13 Dec 1982, P.A. Smith 3981 (BR0000017454506, E, K); between Motswiri and TFC [Tsetse Fly control] road, 18°45.671'S, 23°15.033'E, 966 m, 19 Mar 2003, A. Heath & R. Heath 428 (K); 17 km S of Tsetse Fly control road, 18°45.565"S 23°05.208"E, 964 m, 28 Nov 2004, A. Heath & R. Heath 734 (K); Khwai River floodplain, Moremi Game Reserve, 19°10.505'S, 23°44.176'E, 940 m, 21 Nov 2007, A. Heath & R. Heath 1417 (K); Ngamiland, 100 m S of Samudupe bridge, 20°6'59"S, 23°31'38"E, 938 m, 19 Feb 2008, B. Farrington et al. 486 (K);

Mozambique: Lorenço Marques [Maputo], 7 Dec 1897, B. Schlechter 11643 (BM, BR0000018267990, E, G, LE, P04577163, WAG1103122); Sofala [prov.], Nov 1901, Anonymous 2654 (WAG1103309); Lorenço Marques [Maputo], 21 Mar 1920, Berle 761 (B101143603, BR0000018267983, M); Lorenço Marques [Maputo prov.], between Umbeluzi and Porto Henrique, 20 Nov 1940, A.R. Torre 2090 (K); Gaza prov., Aldeira da Barragem, 20 Nov 1957, L.A.G. Barbosa & F. de Lemos 8222 (K); Gaza prov., Caniçado, 23 Aug 1969, M.F. Correia & A. Marques 1158 (WAG1103272); Gaza prov., Limpopo, Massangena, 21 Jul 1973, M.F. Correia & A. Marques 2979 (K, M, WAG1103326); Gaza prov., Caniçado, 6 Aug 1973, M.F. Correia & A. Marques 3184 (WAG1103323); Lorenço Marques [Maputo prov.], Porto Henrique, Bela Vista, 25 Mar 1975, A. Marques 2670 (WAG1103271); Maputo, 13 Dec 1979, J. de Koning 7754 (BR0000013706678, BM, K);

South Africa: [Limpopo province] Messina [Musina], alt. 2000 ft, Sep 1917, F.A. Rogers 19299a (BM, G); Transvaal [Limpopo prov.], Messina [Musina], 27 May 1927, R.G. Young 18391 (BM); Transvaal prov., Ellisras, 2900 ft, 24 Feb 1954, L.E. Codd 8491 (K, L1698855); Transvaal [Limpopo prov.], Potgietersrust [Mokopane], Doornpoort Farm, 19 Jan 1955, A.D.J. Meeuse 9552 & 9552a (K, M); Transvaal [Limpopo prov.], Waterberg distr., Tamboetie river, Ellisras–Vila Nova road, 7 Jan 1959, A.D.J. Meeuse & R.G. Strey 10446 (BM, BOL217406, BRLU0026260, K, M, Z-000092195); Transvaal [Limpopo prov.], Kruger NP, 31 Jan 1962, H. Schlieben 9321 (W20978); Mpumalanga prov., Kruger NP, 26 Nov 2015, G. Zambatis 1195 (PRE0990262);

Zimbabwe: [Masvingo prov.] Sabi-Lundi Junction District, Chiribira Falls, 6 Jun 1950, Wild 3448 (B101143602, BR0000018267976, K); [Matabeleland South prov.] Gwanda Distr., 700 ft, Nov 1956, R.M. Davies 2185 (K); [Masvingo prov.] Nuanetsi, nr Malipate, 2 May 1961, R.B.Drummond & R.O.B. Rutherford-Smith 7680 (K); [Masvingo prov.] Chiredzi, nr Sabi-Ludi junction, 31 May 1971, S. Mari 1273 (K, LE);

Endemic to Zambezi floristic province (according to Takhtajan 1986). Reported from Okavango region, NE Namibia (Friedrich 1966), but the cited specimen (“Lightfoot 65”) has not been found by us (SAM?).

(Figs 16, 17). Annual, highly branched, with prostrate or ascending stems up to 100 cm long, covered with stellate (sometimes very scattered) and branched hairs, prickles absent. Leaves rosulate, short-lived, and cauline, green or grayish-green turning red at senescence, sparsely to moderately pubescent, rarely glabrous, petiolate (petioles up to 10.0 mm), entire or slightly crisp or scarcely denticulate, ovoid, obovate or oblong-spatulate, 10.0–40.0(45.0) × 3.0–15.0(18.0) mm, acuminate, lateral veins neither recessed adaxially nor prominent abaxially. Flowers in clusters of 8–20, distant or approximated in the upper part of the inflorescence, 8–13 mm in diameter, sessile or with short pedicels up to 3.0(4.0–5.0) mm; buds and closed anthocarp cylindrical. Perianth segments in flowering 3.0–4.2 mm long, in fruiting (3.5)4.0–5.0(5.5) mm long, glabrous or sparsely pubescent, dorsally green or with white margins and ventrally white or mauve; tips ± recurved at fruiting. Stamens (3–4)5(6–8), sometimes with sterile teeth; anthers 0.6–0.9 mm long. Stigmas 3, 0.5–1.0 mm long. Seeds reddish or brown-red, (0.35)0.40–0.50 × 0.25–0.30 mm, colliculate or rarely almost smooth, longitudinal ridges absent.

Mollugo hirta described from South Africa was very rarely accepted in old taxonomic literature, and only cited as a poorly known species (Fenzl 1836). Otherwise, it has been commonly considered a synonym of G. lotoides (e.g., Fenzl 1840; Harvey and Sonder 1860; Just 1879; Adamson 1961; Hedge and Lamond 1975; Gonçalves 1978). We state for the first time that (1) Mollugo hirta must be resurrected in specific rank based on both molecular and morphological studies as G. hirtus, and (2) this species is conspecific with the name G. dahomensis. Specimens of this species were misidentified in collections with various names, particularly G. lotoides, G. lotoides var. virens, G. lotoides × G. oppositifolius, G. spergula, and Mollugo glinoides (both latter names belong to the synonymy of G. oppositifolius). Almeida (1998) referred the Indian plants to G. lotoides subsp. hirtus in the belief that they differ from the European populations (G. lotoides subsp. lotoides), but he did not indicate any differences between them. However, his description of the subspecies rather belongs to G. lotoides based on the number of stamens (>10) and stigmas (5), while G. hirtus has up to eight stamens and three stigmas.

The name G. dahomensis was originally introduced (Chevalier 1938) with a description in French, whereas the nomenclatural code required a description or diagnosis in Latin. To date the name G. dahomensis remains invalidly published because of having been commonly placed in the synonymy of G. lotoides var. virens.

While considering G. hirtus distinct from G. lotoides, we resurrect the name published by Thunberg because it is the only one available for the species as circumscribed in our study.

Glinus hirtus is morphologically very close to G. ononoides Burm.f. described from India (Burman 1768), a species completely forgotten in the past (Dizionario… 1842) and present. The differences between both species are minor and limited to the following characters: (1) the type of G. ononoides has stellate hairs that are mostly localized on the young stem parts and leaves and additionally very short (up to 0.4 mm) prickle-like simple hairs present on the stem. In contrast, G. hirtus has well-expressed stellate pubescence on stem and perianth, and no prickle-like hairs, and (2) the seeds of G. ononoides are smooth, whereas those of G. hirtus are usually colliculate (however, the seeds of the specimens from Senegal, [without date] Perrottet; Botswana, 1982, P.A. Smith; and Somalia, 1893, D. Riva, are smooth). The collections of G. ononoides are very scarce, and the variability of the characters mentioned and the distribution of the species in Asia require further investigations. The morphologically very similar, but phylogenetically distant G. radiatus distributed in South and Central America has shorter anthers (0.35–0.6 mm long) and stylodia (0.2–0.6 mm long) as well as smooth seeds; by contrast, G. hirtus has longer anthers (0.7–0.9 mm) and stylodia (0.5–1.0 mm) and its seeds are usually colliculate (rarely smooth). Also, the distribution areas of both species do not overlap.

River beds, wetlands, damp areas or as a weed, mostly on sandy soils at elevations 0–2400 m a.s.l. Sometimes, G. hirtus is found growing together with G. oppositifolius (collections of Cook 405 & 408 from Nigeria, K!). Observations in Botswana recorded the following associated plants (A. Heath & R. Heath 457, K): Cynodon dactylon, Sida cordifolia, Glinus bainesii, Portulaca oleracea, Cyperus polystachyos, C. compressus, C. longus, Pseudognaphalium luteo-album. Flowers in early and main rains (obs. in Botswana by A. Heath and R. Heath).

(Fig. 18). The species was originally known as Mollugo hirta from South Africa (Thunberg 1794) and not reported as a distinct species from any other African territory. Glinus dahomensis was described from Benin (Chevalier 1938), and later reported from Belgian Congo (DR Congo) (Hauman 1951). Hauman (1951) also noted the presence of the species in other territories of tropical Africa (“Du Dahomey au Transvaal”). However, he probably was not sure of that and did not reidentify the specimens from any other countries. We confirm that the range of G. hirtus is not restricted to Benin and DR Congo, but the species is distributed in all sub-Saharan Africa and seems to be a common weed in many regions according to the collector’s observations. Glinus hirtus has not been previously reported for almost any Sub-Saharan countries (e.g., Adamson 1961; Jeffrey 1961; Berhaut 1967; Gonçalves 1970, 1978; Sita and Moutsambote 1988; Barry and Celles 1991; Mapaura and Timberlake 2004; Phiri 2005; Setshogo 2005; Hassan et al. 2005; Sosef et al. 2006; Germishuizen et al. 2006; Darbyshire et al. 2015). In some West African checklists and manuals it was confused with the American G. radiatus (Berhaut 1967, 1979; Boudet et al. 1986; Akoegninou et al. 2006; Lisowski 2009; Thiombiano et al. 2012; Brundu and Camarda 2013; Schmidt et al. 2017). Only a few specimens from Burkina Faso, DR Congo, and Nigeria were correctly identified as G. dahomensis.

Angola (new records): [Moxico prov.] Chizanda, 23 Sep 1899, H. Baum 135 (BM, BR0000018269482, K, M); [Malanje prov.] Malange [Malanje], 21 Sep 1932, R.G.N. Young 846 (BM); Huila [prov.], Lubango, 11 Dec 1955, E.J. Mendes 1083 (BM, LISC032007); Luanda, Namuculungo, 1100 m, 13 Nov 1957, J.B. Teixeira 3185 (LISC032013); Huila prov., Canguelas, Vila Artur de Paiva, 1450 m, 30 Dec 1959, E.J. Mendes 1879 (M); Benguela prov., Ganda, 30 Jun 1960, J.M. Teixeira & A.M. Andrade 4988 (LISC032014); Cuanza Sul prov., Uaco-Cungo, 2 Nov 1961, J.M. Teixeira & J.M. Figueira 5956 (LISC032015); [Cunene prov.] Mucope, Tchica [river], 9 Nov 1963, C. Henriques 217 (BM, K, LISC032005); Huila prov., Gambos, 18 Nov 1970, R.M. Santos & E. Barroso 2874 (LISC032010); Huila prov., Quipungo, 1200 m, [without date] J.M. Teixeira 12580 (LISC032004);

Benin: see type of G. dahomensis; Atakora [dept.], Toukountouna, 12 Apr 1999, A. Akoegninou et al. 2440 (WAG0235864); Atakora [dept.], Kérou, 16 Apr 1999, A. Akoegninou et al. 2568 (WAG0235868); [Donga dept.] Bassila, 460 m, 30 May 2001, A. Akoegninou 4810 (WAG0235872);

Botswana (new records): Northern distr., Nxauna Pan, 1 Jan 1973, P.A. Smith 318 (K); [North-West distr.] nr Tsau–XaiXai road, 19°55'2"S, 21°24'4"E, 21 Apr 1982, P.A. Smith 3847 (BR0000018269659; PRE0671232, PRE0520975); [Ngamiland distr.] Ngami Lake, 13 Dec 1982, P.A. Smith 3980 (BR0000018269666, E, G, PRE0671267); Ngamiland distr., between Caecae and Gwihaba Hills, 1000 m, 15 Mar 1987, D.Anonymous Long & D.A.H. Rae 300 (E, K); Zibalianja lagoon, 18°34.728'S, 23°32.145'E, 957 m, 23 Jan 2004, A. Heath & R. Heath 457 (BACH, K);

Burkina Faso (new records): Kompienga [prov.], 40 km E of Tindangou, 2 May 2003, L. Sanou & M. van Slageren 1352 (K); Houet [prov.], 18 km NW of Bobo Dioulasso, 12 May 2003, L. Sanou & M. van Slageren 1390 (K);

Burundi (new records): [Karuzi prov.] Nyabibuye, 4500 ft, 15 Nov 1960, R. Tanner 5602 (K, WAG1103206); [Ruyigi prov.] Ruyigi, 2°59'S, 30°28'E, alt. 1450 m, 27 Sep 1978, M. Reekmans 7098 (K);

Cameroon (new records): Southwest region, Mamfe distr., 15 Mar 1953, C.F.A. Onochie et al. 30889 (K); [Centre Province region] 62 km SE of Bafia, 27 Mar 1963, J. Raynal & A. Raynal 10537 (P0456668); [North region] nr Garoua, Benoue river, 18 Dec 1964, W.J.J.O. de Wilde et al. 4959 (WAG0180494); [Litoral region] plaine des Mbo, 7 km E of km 25 Melong–Dechang (Dschang) road, 5°11'24"N, 10°1'12"E, alt. 700 m, 14 Apr 1972, A.J.M. Leeuwenberg & C.C. Berg 9607 (BR0000018268621, K, P04576510, WAG0185131); North region, Bénoué NP, 7 May 1974, C. Geerling & J. Néné 4705 (BR0000018268591, WAG0330521); [Centre region] river Sanaga, bridge nr Nkol Ngok, 4°10'N, 11°01'E, alt. ~300 m, 15 Feb 1978, J. Lowe 17675 (K);

Central African Republic (new records): Kémo [pref.], 17 Feb 1892, Anonymous 691 (P04577135); Ouaka pref., Bambari, 7 May 1927, R.P. Tisserant 2180 (p04577138); [Haute-Kotto pref.] 300 m NW of Gounda, 9°18'N, 21°12'E, alt. 450 m, 28 Jul 1982, J.M. Fay 2763 (K001394828);

Chad (new record): [Logone Occidental region] Kim [Krim], 18 May 1969, Anonymous Fotius 1458 (P04576514);

DR Congo (selected): [nr Kinshasa] Stanley Pool [Pool Malebo], 1888, F. Hens 29 (BR0000018268034); [Kongo Central prov.] Kisantu, 1900, J. Gillet 1386 (BR000018268027); Kongo Central prov.] Kisantu, 1903, J. Gillet 3029 (BR000018268010); [Kasai prov.] Zambi vill., 27 Jun 1915, Bequaert 7885 (BR000018268003); [Kwilu prov.] Wombali [Bandundu], Oct 1915, H. Vanderyst 2316 (BR0000018268171); Haut-Katanga prov., Kafubu, 18 Nov 1927, Quarré 804 (BR0000018268386); [Mongala prov.] Bumba, 19 Feb 1931, R. Letouzey & J.F. Villiers 10386 (K, P04576657, WAG0185129); [Haut-Uele prov.] Garamba NP, 20 Feb 1950, H. de Saeger 147 (BR0000018268232); Orientale prov., Dungu, 2 Apr 1951, H. de Saeger 1084 (K); Orientale prov. [Haut-Uele prov.] Dungu, Garamba NP, 28 Mar 1951, H. de Saeger 1119 (K); [Lualaba prov.] Dikuluwe, 21 Aug 1956, J. Brynaert 520 (BR0000018268324, K); [Haut-Katanga prov.] Mitwaba, 2 Nov 1956, J. Brynaert 541 (BR0000018268317, K, LE, L1698856, M, W13591); [Tshopo prov.] Yangambi, 5 Feb 1958, R. Devred 2925 (WAG0185147); [Kasai-Oriental prov.] Katanda Territory, 17 Sep 1959, L. Liben 3701 (BR0000017455121); [Maniema prov.] Kabambare Territory, 10 Aug 1959, 570 m, A. Leonard 5736 (BR0000018268201, WAG0185149); [Maniema prov.] Kamisuku, 700 m, 20 Aug 1959, A. Leonard 6034 (BR0000018268218, P04576678, WAG0185148); Yangambi prov., Bangi, Ile Yangambi, 7 Sep 1959, P. Bamps 700 (BR0000018268188, K); [Haut-Lomami prov.] Bukama, Lualaba river, 570 m, 2 Oct 1970, M. Lukuesa 758 (BR0000018268355, WAG0185146); Kongo-Central prov., Kasangulu, Sabuka, 6 Apr 1972, C. Evrard 6929 (BR0000018269673); Haut-Katanga prov., Kumanua, 1150 m, 18 Dec 1975, F. Malaisse 8869 (BR0000018268379); [Eastern prov.] Waine-Rukula, 22 Oct 1978, J. Lejoly 4180 (BR0000017455336); [nr Kinshasa] Inkisi river, 17 Oct 2007, Nsimundele 2060 (BR0000000530230);

Ethiopia (new records): [Amhara region] Begember prov., [years] 1863–1868, Schimper 1340 (K); [Oromia region] Jimma, Apr 1957, Anonymous S2 (K); Godjam prov. [Amhara region], Bahir Dar, 1800 m, Jun 1968, O. Sebald 2166 & 2255 (M, STU);

Gabon (new records): Moyen-Ogooué prov., Lambaréné, 17 Oct 2012, Boupoya 760 (BRLU0026277); Ogooué-Maritime prov., Wonga-Wongué Reserve, 20 Oct 2014, A. Boupoya & al. 1087 (BRLU0026275); [Moyen-Ogooué prov.] Onangué Lake, 35 km SE of Lambaréné, 22 Oct 2014, E. Bidault et al. 1820 (BR0000016174696, BRLU0026276, P00854718);

Gambia (new record): [North Bank Division] Albreda, 1839, Haudelot 100 (G);

Ghana: [Oti Region], on road to Oti from Kete Krachi, 18 May 1952, J.K. Morton 7294 (K); [Upper West region] Tumu, edge of Dam, 25 May 1952, J.K. Morton 7554 (K, WAG0185143); [North East region] Gambaga, 26 Dec 1954, J.K. Morton A1378 (K);

Guinea (new records): [Kankan region] Yri Kiri, 19 May 1955, Anonymous Roberty 18001 (G); [Kankan region] Kankan, Bordo, 26 Apr 1956, J.-Anonymous Adam 12119 (P00695349); [Nzérékoré / Kankan Regions] Simandou Range, Oueléba swamp, alt. 1270 m, 19 Oct 2008, I. Darbyshire 495 (K000580882); [Nzérékoré / Kankan Regions] Simandou Range, Canga East, May 2009, P.K. Haba 579 (K000024353, P00990780, WAG1476650);

Guinea-Bissau (new record): [Bafatá Region], Bambadinca, 11 Jun 1945, Anonymous 2065 (K, WAG0185158);

Ivory Coast (new records): [Vallée du Bandama distr.] Fétékro, 18 Jan 1947, Anonymous Roberty 6915 (G); [Bouna dept.] Bouna, 1 Dec 1986, P. Poilecot 3833 (G40376);

Kenya (new records): [Nairobi] Nairobi Dam, 31 Jan 1961, C. Teesdale 26 (K); Nairobi National Park, 1525 m, 7 Oct 1977, J.B. Gillett & W.T. Staam 21582 (BM, K); [Eastern prov.] 36 km from Embu town, 14 Nov 1979, AnonymousW. Gatheri et al. 79/1 (K);

Malawi (new records): [Northern region] nr Karonga, 1893–1894, AnonymousF. Scott-Elliot 8403 (BM, K); Zomba, 2500–3500 ft, A. Whyte s.n. (G, P04576503); Nyasaland [without exact location], 1895, J.Buchanan 15 (BM, BR0000018269611, E); [Central region] Dedza Distr., Chongoni Forest reserve, 18 Sep 1967, A.J. Salubeni 832 (K); [Central region] Kasungu distr., Kasungu NP, 12 Dec 1970, A.J. Hall-Martin 1283 (K); [Central region] Kasungu distr., Kasungu NP, 1000 m, 23 Dec 1976, J. Pawek 12032 (K, WAG1103230); [Southern region] Liwonde NP, 9 Oct 1978, A.J. Salubeni & E. Tawakali 2339 (WAG1103225); Central Region, Mchinji distr., 21 Nov 1983, A.J. Salubeni & I.H. Patel 3484 (BOL217407); [Central region] Lilongwe Distr., Nature Sanctuary, 8 Nov 1984, I.H. Patel & al. 1656 (K); Central Region, Nkhotakota distr., 13 May 1986, I.H. Patel & R.B. Kwatha 3184 (BRLU0026262); Northern region, Chitipa, 9°58'51"S, 33°25'37"E, 1528 m, 18 Dec 2007, H.T. Chapama & al. 788 (K);

Mali (new records): [Koulikoro region] Koulikoro, 11 May 1912, Vuillet 609 (P04577141); [Inner Niger Delta region] Mopti, 24 Apr 1932, J. Rogeon 76 (P04577146); [Gao region] Bamba, 1927, O. Hagerup 327 (BR0000018268447, K, P04576677); [Gao region] Gao, 21 Jun 1936, M. de Wailly 5070 (P04577142); [nr Bamako] Niger river, Sotuba, 4 Jul 1973, D. N’Golo 1291 (WAG0319754); [Sikasso region] Sikasso, Yanfolila, 7 Jul 2009, L. Sanou et al. 624 (K);

Mozambique (new records): [Nampula prov.] Nampula, Mutivasi river, 3 Nov 1942, F.A. Mendonça 1217 (K); [Zambezia prov.] Boroma, 4 miles from Zambesi river, 25 Jul 1950, N.C. Chase 2788 (BM); [Sofala prov.] Gorongosa distr., 300 ft, 26 Sep 1953, N.C. Chase 5087 (BM); Sofala prov., Gorongoza Game Reserve, 25 Aug 1958, N.C. Chase 6980 & 6981 (BR0000018269185, FT0007098, K); Sofala prov., Gorongosa NP, 3 Nov 1963, A.R. Torre & J. Paiva 9006 (BR0000018269161, M); [Sofala prov.] Beira, Jul 1970, K.L. Tinley 1936 (BR0000018269642); Sofala prov., Gorongosa NP, Jul 1970, K.L. Tinley 1938 (K); Gaza prov., Caniçado, 17 Nov 1970, M.F. Correia 2018 (WAG1103222); Niassa prov., Lugenda river, Marrupa to Mecula, 11 Aug 1981, P.C.M. Jansen et al. 225 (K, WAG0312529); Sofala prov., Gorongosa NP, 18°55'4.8"S, 34°30'38.04"E, 9 Nov 2007, P. Ballings & B. Wursten 944 & 958 (BR581135 & BR0000018269635);

Namibia (new records): [Otjozondjupa Region] Okahandja, Nov 1906, K. Dinter 301 (BM, BR0000018269222, E, G, K, P04577101, U139844, WU, Z-000092198); [Khomas region] Windhoek, 21 Dec 1929, C.E. Moss 18055 (BM); [Khomas region] Khomas Highland, Friedenau Farm, 2000 m, 6 Apr 1939, Anonymous Gassner 124 (M); [Erongo region] Swakopmund distr., Goabeb Research Station, 9 Oct 1963, C. Koch s.n. (K, PRE0403669); [Hardap region] 30 miles E of Maltahöhe, 4400 ft, 15 Feb 1950, E.C.Macdonald 369 (BM); [Kavango-East region] Rundu [between Kapuko & Sambiu], 22 Dec 1955, B. de Winter 4035 (K, M, PRE0403672); [Khomas region] Windhoek, 1575 m, 29 Apr 1961, R. Seydel 2819 (B101143626, G, L1693130); [Erongo region] Swakopmund, mouth of Swakop river, 11 Feb 1962, R. Seydel 3041 (B101143623, M); [Khomas region] Windhoek, Swakop river, 2000 m, 20 Feb 1965, R. Seydel 4194 (B101143629, B101143630, K);

Niger (new record): [Tillabéri region] Kouré, 8 Mar 1897, Anonymous Roberty 2556 (P04577117);

Nigeria (new records): [Benue State] Abinsi, 29 May 1912, J.M. Dalziel 715 (K); [Jigawa State] Takara, 3200 ft, 4 May 1921, H.V. Lely 101 (K); Adamawa State, Jimeta (Yola), 29 Dec 1957, F.N. Hepper 1621 (K); [Niger State] Shagunu, 10 km N of Bussa, 25 Jul 1965, C.D.K. Cook 405 (K); Niger State, Bida distr., 1 Mar 1968, B.O. Daramola & A. Binuyo 61930 (K); [Taraba State] Sardauna area, Bissaula, 18 Dec 1968, B.O. Daramola 62328 (K); [Taraba State] Gashaka distr., Gashaka river, 1100 ft, 25 Mar 1970, Z.O. Gbile & B.O. Daramola 23888 (WAG0185150); Bauchi State, Yankari Game Reserve, 24 Mar 1971, C. Geerling 3486 (B100480213, BR0000017456135, WAG0083282); Gombe State, Dadin Kowa, Gongola river, 2 May 1972, Gbile et al. 65453 (K); Adamawa State, Yola distr., 6 May 1972, Gbile et al. 1303 (K); Enugu State, Amechi vill., 21 Feb 1973, Latilo & Oguntayo 67620 (K, WAG0185136); [Kogi State] Koton Karfe, Niger river, 22 Mar 1973, Eimunjeze et al. 70400 (K); Niger State, Kafin Koro, 23 May 1973, Eimunjeze et al. 66496 (K);

Republic of the Congo (new record): [Bouenza dept.] Jacob [Nkayi], 27 Sep 1969, Y. Attims 255 (WAG00034942);

Rwanda (new records): Mutara area, 1400 m a.s.l., 3 Oct 1956, Anonymous Troupin 2810 (BR0000017455824, K); [Eastern prov.] Nyagatare, 1450 m, 27 Jan 1958, Anonymous Michel 5077 (BR0000017455817);

Senegal (new records): [without exact location and date] Perrottet 373 (G, P04577123); Wab, 20 Dec 1824, Perrottet 51 (P04577122); [Waalo Region] N’ghiau, [without date] Lepier s.n. (G); Oussodou, 2 Jan 1954, R.P. Berhaut 4330 (P04577147); [Ziguinchor Region] Bignona, 22 May 1957, J.-Anonymous Adam 13656 (P04577113); Niokolo Koba NP, Feb 1960, J.-Anonymous Adam 17485 (P04577114); Basse Casamance [Ziguinchor], 11 Jul 1963, R.P. Berhaut 6084 (BR7000919, P04577119); Basse Casamance [Ziguinchor], 29 Jul 1977, C. Vanden Berghen 1915 (BR0000018268461, WAG0106332);

Sierra Leone (new records): [Northern prov.] Samaia, 10 May 1914, N.W. Thomas 245 (K); [Northern prov.] Mange, 21 Feb 1949, F.C. Deighton 4978 (K); [Northern prov.] Yifin, 23 Mar 1964, Norton & Gladhill 994 (K, WAG0185139); [Northern prov.] Koinadugu distr., SE of Fadugu, 21 May 2014, X.M. van der Burgt 1854 (K);

Somalia (new record): [Gedo prov.] nr Ganane, Mar 1893, D. Riva 736 (G, RO);

South Africa: [Gauteng prov.] Pretoria, Apies river, [without date] ex herb. J. Burtt Davy s.n. (K); [Gauteng prov.] Pretoria distr., Apies river, [without date] Zeyher 612 (K); [Western Cape] Clanwillam, Feb [no year] Zeyher & Ecklon 1818 (P04577109); [Western Cape] Clanvillam, secus fluvium Olifantsrivier ad villam Brackfontein, Jun 1847 Zeyher 612 (P04577110, P04577124); Cape of Good Hope, 1847, C.-L. Zeyher 612 (FR-0132679, G, LE, P04577110); Northern Cape, Henkries, 30 Nov 1897, B. Schlechter s.n. (BR0000018269291, G, LE, L1693128, P04576623, WAG1103197, Z-000092196); [Limpopo prov.] Musina, 2000 ft, [without date] F.A. Rogers 22240 (G); [KwaZulu-Natal prov.] Durban, Sep 1904, M. Wood s.n. (L1693126); [Limpopo province] Messina [Musina], Nov 1917, Moss & Rogers 57 (BM, K); [Mpumalanga prov.] Klipfontein, Nov 1937, J.P.H. Acocks 5145 (K); [Transvaal prov.] Zoutpansberg, Kruger Nat. Park, 800 ft, 2 Nov 1948, O’Dyer 4630 (K); [Northern Cape] Lokenburg, 2100 ft, 16 Mar 1957, J.P.H.Acocks 19196 (K, M, PRE0403696, PRE0403696-2); [Northern Cape prov.] Kimberley, Rooipoort, 3450 ft, 1 Dec 1958, O.A. Leistner 1210 (K); Cape Prov., [Northern Cape prov.] Warrenton, [without date] J.P.H.Acocks 15916 (K); [KwaZulu-Natal prov.] Ubombo Distr., Mkuzi Gane reserve, 9 Dec 1959, C.J.Ward 3353 (K, M); Transvaal prov., Kruger NP, 350 m a.s.l., 27 Jan 1982, J. Lambinon & M. Reekmans 82 (BR0000018269246, MSB-129442, PRE0626149); [Limpopo prov.] Giyani distr., 18 Oct 1988, S. Venter 13049 (PRE0810982);

South Sudan (new record): [Equatoria Region] Zande, 6 Mar 1940, J.W.AnonymousWyld 781 (BM);

Tanzania (new records): [Ruvuma region] 7 km W of Songea, 990 m, 18 Jan 1956, E. Milne-Redhead & P. Taylor 8355 (BR0000018269017, K); [Rukwa region] Sundu Lake, 1500 m, 13 Dec 1958, H.M. Richards 10316 (K); Kigoma region, Mugombasi, 1 Sep 1959, R.M. Harley 9495 (K); [Kagera region] Ngara, 4500 ft, 6 Sep 1960, R. Tanner 5126 (BR0000017454308, K, WAG1103204); [Singida region] Ruaha NP, Great Ruaha river, 840 m, 19 Jan 1966, M. Richards 21007 (K); Arusha region, Ngurdoto Crater NP, 22 Mar 1966, P.J. Greenway & Kanuri 12474 (BR0000017454285, K); Arusha region, Arusha NP, 1524 m, Nov 1969, M. Richards 24683 (K, M); [Singida region] Ruaha NP, 820 m, 12 Oct 1970, A. Bjørnstad 601 (K); [Arusha region] Ngorongoro Conservation area, 2200 m, 30 Dec 1988, T. Pocs & S. Chuwa 88308 (K); Mbeya region, 19 Dec 1989, J. Lovett et al. 3775 (L0717072, P04576671); [Kilimanjaro region] Kilimanjaro, 1220 m, 2 May 1994, J.M. Grimshaw s.n. (K);

Uganda (new record): [Western region] Kiryadongo, 3200 ft, Mar 1943, Anonymous 1338 (K);

Zambia (new records): [North-Western prov.] Mwinilunga distr., 6 Nov 1937, E. Milne-Redhead 3115 (BR0000017454414, K); [Southern prov.] Mapanza, 20 Sep 1953, E.A. Robinson 324 (BR0000018269109, K); [Copperbelt prov.] Ndola, 23 Jun 1955, D.B. Fanshawe 2343 (B101143625, BR0000018269604, K); [Lusaka prov.] S of Lusaka, Kafue river, 3000 ft, 11 Dec 1955, E.B. Best 121 (K); [Eastern prov.] Lundazi, Tigone Dam, 1200 m, 19 Nov 1958, N.K.B. Robson 655 (BM, K); [Southern province] Choma, 25 Oct 1958, E.A. Robinson 2900 (M); [Eastern prov.] stream beside Chadiza turn-off, Nsadzu to Fort Jameson [Chipata], 900 m, 25 Nov 1958, N.K.B. Robson 703 (BM, BR0000017454438, K); [Northern prov.] Kali Dambo, [without date], H.M. Richards 4091 (K); [Central prov.] Broken Hill [Kabwe], 14 Oct 1959, J.M. Mutimushi 74 (K); [Eastern prov.] Chimutengo/Petauke, 3 Oct 1966, J.M. Mutimushi 1542 (K); Lusaka prov., Chitendabunga, 1200 m, 13 Nov 1993, M.Anonymous Bingham 9795 (WAG1103214);

Zimbabwe (new records): Bulawayo, May 1898, R.F. Rand 331 (BM); Salisbury [Harare], Sep 1919, Eyles 1806 (K); [Matabeleland North prov.] Victoria Falls, Nov 1933, A. Meebold 11992 (M); [Matabeleland South prov.] Beitbridge, Limpopo river, 21 Dec 1935, Smuts & Gillett 3108 (PRE0800058); [North-Western prov.] Salisbury [Harare], Makabusi river, 4500 ft, 24 Oct 1945, H. Wild 269A (K); [Matabeleland South prov.] Bulalimamangwe distr., Tegwani, 8 Jan 1946, F.W.J. McKosh 14297 (K); [Manikaland prov.] nr Mutare, 3500 ft, 29 Dec 1946, B.S. Fisher 16533 (K); Nuanetsi, 1850 ft, Nov 1955, R. Davies 1610 (K); Bulawayo, 4800 ft, Dec 1956, O.B. Miller 4000 (BR0000018269628, K); [Mashonaland West prov.] Urungwe distr., Zwipani Vlei, 30 Nov 1957, R. Goodier 425 (K); [Matabeleland South prov.] Nuanetsi, 2050 ft, 6 May 1958, R.B. Drummond 5599 (K); [Matabeleland South prov.] Gwanda Distr., nr Chikwarakwara, 12 May 1958, R.B. Drummond 5789 (K); [Mashonaland Central prov.] Sipolilo, Angwa river, 3 Jun 1965, M.Anonymous Bingham 1548 (BR0000018269550); [Midlands prov.] Owelo [Gweru], 4200 ft, 30 Dec 1965, B.K. Simon 587 (K); [Midlands prov.] Kwekwe distr., Sable Park, 26 Feb 1976, J.M. Stephens 340 (M, WAG1103229);

Africa (Sub-Saharan Africa; North Africa: Egypt [WU!]; Madagascar: G! K!).

(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.

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).

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.

(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);

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.

Annual, branched from the base, with numerous prostrate stems often forming mats up to 1 m in diameter; young parts of the stems covered with simple crispate hairs usually arranged along one line; prickles absent or unnoticeable with the naked eye. Leaves rosulate, short-lived, and cauline, green, glabrous or puberulent, shortly petiolate (petioles 3–10 mm long), entire or coarsely denticulate, oblong, ovate, obovate or narrowly obovate, (10)15–45(55) mm × 3–15(20) mm, apically shortly acuminate, lateral veins neither recessed adaxially nor prominent abaxially. Flower clusters interrupted, consisting of (1)2–10 flowers, rarely more; flowers usually with unequal pedicels 7.0–20.0 mm long, or sometimes (sub)sessile; buds and closed anthocarp of cylindrical shape. Perianth segments in flowering (2.7)3.0–3.5 mm long, in fruiting (3.5)4.0–5.0(5.5) mm long, glabrous or sparsely pubescent, dorsally green or pink with white margins and ventrally white or pinkish, sometimes turning red at senescence; petaloids usually absent. Stamens usually 5, rarely 4 or 6–7; anthers (03.–0.5)0.6–0.8(1.0) mm long. Stigmas 3, 0.3–0.6 mm long. Seeds reddish or brown-red, rarely yellow-brown, 0.30–0.50 × (0.25)0.3–0.4 mm, colliculate or rarely smooth, longitudinal ridges absent.

Glinus oppositifolius is sometimes confused with Gisekia pharnaceoides L. (Gisekiaceae). The most remarkable characters of Gisekia are the leaves and perianth with white striae (formed by cells with raphides), and its capsules are divided into 5–15 mericarps each containing one exarillate seed (Gilbert 1993b). In contrast to Gisekia, Glinus species do not have easily visible raphides in any part of the plant and their capsules are multi-seeded with arillate seeds.

Sands, river banks or as a weed; altitude up to 1600 m a.s.l. The most common variety in Sub-Saharan Africa. The associated plants found in Botswana (A. Heath & R. Heath 456, K): Cynodon dactylon, Sida cordifolia, Glinus bainesii, G. hirtus, Portulaca oleracea, Cyperus polystachyos, C. compressus, C. longus, Pseudognaphalium luteo-album (formerly known as Helichrysum luteo-album). Flowers mostly during the main rains (in southern Africa).

(Fig. 24). Angola: [Bengo prov.] Barra do Bengo distr., Sagoa, Dec 1853, F. Welwitsch 1265 (BM); [Luanda prov.] between Camama and Calumbo, Jul 1854, F. Welwitsch 1110 (BM); Luanda distr., between 8 and 9° S, Jan 1858, F. Welwitsch 2409 (BM, LE); [Bengo prov.] Barra do Dande distr., Dande river, Sep 1858, F. Welwitsch 1112 (BM); [Zaire prov.] Tombe, 1908, H.A. Junod 2886 (BM); Luanda, Muceque, Mar 1914, J. Gossweiler 5880 (COI00070554); Luanda, 1918, K. Müller 1109 (G); [Kwanza Sul / Bengo provinces] Catemba to Muceque, 60 m, 30 Sep 1935, J. Gossweiler 10405 (BM); [Bengo prov.] Calemba, 60 m, 30 Sep 1935, J. Gossweiler 10405 (BM, COI00070555);

Benin: Porto Novo, 6 Mar 1910, A. Chevalier 23314 (P04577130); Cotonou, 19 Jan 1962, D. Froment 1141 (BR0000018269932); Ouémé [dept.], Bonou, 12 Mar 1998, P. Houngnon 5608 (BR0000018269949, K); [Collines dept.] Dassa, 4 Nov 1998, V. Adjakidjè et al. 2189 (BR0000018269956, K, WAG0095242); Zou [dept.], Djidja, 11 Feb 1999, J.-P. Essou et al. 1251 (WAG0235854); [Kouffo dept.] Klouékanmé, 15 Feb 1999, J.-P. Essou et al. 1340 (WAG0235856); Borgou dept.], Tchaourou, 7 Apr 1999, A. Akoegninou et al. 2258 (WAG0235860); [Alibori dept.] Malanville, 28 Apr 1999, P. Houngnon et al. 6545 (WAG0235849); Mono dept., Athiémé, 3 Jul 1999, N. Sokpon et al. 838 (WAG0235853); Zou dept., Cové, 14 Jul 1999, P. Houngnon 6798 (WAG0235850); [Donga dept.] Bassila, 20 Apr 2000, P. Houngnon 7634 (WAG0235851); [Collines dept.] Savalou, 9 Jun 2000, V. Adjakidjè 3595 (WAG0235859); Atakora dept., Natitingou, 28 Jun 2000, N. Sokpon 1610 (WAG0317484); Mono dept., Comé, 14 Aug 2000, V. Adjakidjè 3750 (WAG0235874);

Botswana: Northern distr., Mutsoi, 9 Nov 1967, F.L. Lambrecht 420 (K, PRE0403655); Ngamiland distr., Okavango swamp, 25 Feb 1973, P.A. Smith 449 (K, PRE0521257); [Central distr.] Boteti river, 20°23'75"S, 24°31'4"E, 15 Feb 1980, P.A. Smith 3031 (PRE0671278); [North-West distr.] Okavango delta, Maun, 15 May 1984, P.A. Smith 4443 (BR0000017454513, K); [Ngamiland distr.] Zibalianja lagoon, 18°34.7'S, 23°32.15'E, 957 m, 23 Jan 2004, A. Heath & R. Heath 456 (BACH, K);

Burkina Faso: Oudalan prov., Oursi, 22 Sep 1995, S. Kahlheber 145 (FR-0014454); Oudalan prov., Yomboli, 24 Jul 1999, J. Müller 86 (FR-0014433); Oudalan prov., Darkoye, 8 Aug 1999, J. Müller 142-c (FR-0024423); Kompienga prov., 11°09'39"N, 0°37'24"E, 203 m, M. Schmidt 720 (FR-0021934); Ouagadougou, 8 Jun 1987, J. Lejoly 87/020 (BRLU0026274);

Burundi: Bujumbura, 800 m, 29 Oct 1968, J. Lewalle 3076 (BR0000017455862); Bujumbura, 800 m, 28 Sep 1971, J. Lewalle 6144 (G, WAG0104163); Bubanza prov., Randa, 950 m, 25 Sep 1976, M. Reekmans 5325 (BR0000017455909, H1235324, K, WAG0185122);

Cameroon: [Southwest region] Mamfe, on the sandbanks by the Cross river, 15 Mar 1953, C.F.A. Onochie et al. 30889 (K); [Littoral region] nr Masok, Ouem river, 4 Apr 1965, A.J.M. Leeuwenberg 5378 (BR0000018269987, K, P04577247, WAG0185118); [Littoral prov.] 45 km SE of Yabassi, 18 Jan 1972, R. Letouzey 11022 (BR0000018270006, P04577207, WAG0185121); [Littoral prov.] Mbombe river, 20 km W of Yabassi, 1 Mar 1976, R. Letouzey 14779 (BR0000018269994, K, P04577210, WAG0185120);

Chad: [Chari-Baguimi region] Mailao, 1 May 1966, Stauch 23 (P04577261); [Chari-Baguimi region] between Djimtilo and Lake Chad, 31 Jan 1968, J. Léonard 4445 (BR0000017456012, G, K, M, P04577268, PE01696734, WAG1103324); [Chari-Baguimi region] Ouazkaga, 27 May 1971, Anonymous Fotius 1938 (P04577255); Sarh, 7 Jun 1975, Palayer 605 (P04577265);

DR Congo (selected): [Kwilu prov.] Wombali, Nov 1910, Vanderyst s.n. (BR0000013706029); [Province of Équateur] Bokala, Anonymous 4777 (BM); [Congo Central prov.] Boma, Congo river, 27 Sep 1913, Bequaert 804 (BR0000017454537); [Kongo Central prov.] Matadi, Nov 1913, Verschueren 920 (BR0000017455060); [Orientale prov.] Laga, 10 May 1914, J. Gossweiler 5880 (BM); Leopoldville [Kinshasa], Jul 1915, L. Achten 17 (BR0000017454667); Bas-Congo [Congo Central prov.], 1921, J. Claessens 27 (BR0000017454568); [Shopo province] Stanleyville [Kisangani] 25 Jan 1926, W. Robyns 1418 (K); [Province of Équateur] Bamania, 1930, J. Lebrun 884 (K); [Congo Central prov.] Boma, 4 Nov 1930, Vanderyst 27235 (BR0000017454612); [Orientale prov., Tshopo distr.] Yangambi, 6 Jun & 12 Jul 1938, J. Louis 9711 & 10313 (BM, BRLU0026266, K, P04577193, U1398742); [Province of Équateur] Eala, 5 Aug 1946, J. Léonard 240 (ALCB012594 – image seen! FT0007107, K, P04577194); [Kongo Central prov.] Vista [Nsiamfumu], 6 Nov 1947, L. Toussaint 27 (BR0000017454599); Katanga prov., Mitwaba, Nov 1948, AnonymousF. de Witte 4816 (K); [Tswhopo prov.] Basoko, Jun 1949, R. Germain 4959 (BR0000017455268); [Haut-Uele prov.] Uele, 3 Mar 1952, Anonymous Troupin 282 (K, WAG0185123); [Kwango prov.] Popokabaka, 23 Sep 1952, H. Callens 3707 (BR0000017544780, WAG1103266); [Orientale prov.] Albert Lake, Semliki, 5 Mar 1954, D. van der Ben 1184 (BR0000017455664, K); [Sud Ubangi prov.] Zongo, 13 Jul 1957, C. Evrard 2561 (BR0000017455602); Coquille Territory, Indjolo, 3 Oct 1957, C. Evrard 2611 (BR0000017455237); [Haut-Katanga prov.] Lupoto, 27 Nov 1957, A. Schmitz 6031 (BR0000017455763); [Sankuru prov.] [Province of Équateur] Bolomba, 3 Sep 1958, C. Evrard 4948 (BR0000017455251); [Tshopo prov.] Yangambi, 7 Sep 1959, P. Bamps 701 (BR0000017455169; WAG0185116); [Bas-Congo prov.] Seke-Banza, Isangila, 22 Sep 1959, P. Compere 438 (BR00000174554797); [Kongo Central prov.] Kasangulu, Ngombe, 16 Jul 1964, L. Pauwels 4590 (BR0000017454988); Katanga prov., Kilwa, 900 m a.s.l., 31 Dec 1965, J.-J. Symoens 11996 (K); [Haut-Lomami prov.] Lualaba river, Bukama, 2 Oct 1970, M. Lukuesa 740 (BR0000017455756; WAG0185117); Haut-Katanga prov., Kilwa, Moero Lake, 16 Oct 1970, S. Lisowski 61915 (BR0000017455725); [Tshopo prov.] Kisangani, 5 Nov 1977, J. Lejoly 2075 (BRLU0026269); [Tshopo prov.] Ubundu, 12 Jul 1981, Ndjele 427 (BRLU0026273); [Kongo Central prov.] Luozi, 28 Sep 1986, H. Breyne 5215 (BR0000017454759); [Kwilu prov.] Kikwit, 12 Aug 1991, B. Masens 1013 (BR0000017455084, K, WAG0405337); [Tshopo prov.] Lokutu, 7 Nov 2004, Q. Luke et al. 106292 (K); Moenge, Itimbiri river, 360 m, 17 May 2010, Boyekoli Ebali Congo Exp. 501 (BR0000000571554);

Ethiopia: [Southern Nations, Nationalities and Peoples’ region, Keffa zone] Gogeb river, Bonga road, 1300 m, 25 Feb 1966, W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes 10194 (BR0000018270037, K, WAG1103268); [Oromia region] Illubabor zone, Alwero river, Abobo, 630 m, 20 Apr 1982, I. Friis & al. 2475 (K);

Gabon: [Ogooué-Maritime prov.] Port Gentil, 12 Sep 1968, F.J. Breteler & R.A. van Raalte 5526 (WAG0185127); N of Libreville, 13 Aug 1992, J. Dibata 1055 (WAG0070773);

All other specimens identified as Glinus oppositifolius indeed belong to G. hirtus, Gisekia pharnaceoides (Gisekiaceae) and Polycarpaea sp. (Caryophyllaceae).

Ghana: Volta river, 27 Mar 1922, J.M. Dalziel 106 (K, M); [nr Accra] Pokuase, Apr 1931, F.R. Irvine 1589 (E, K); [Ashanti region] Fumso, 24 Mar 1950, Anonymous 543 (K); [Western region] Princes Town, 11 Mar 1952, J.K. Morton 6607 (K, WAG1103299); [Oti region] Kete-Krachi, 19 May 1952, J.K. Morton 7288 (K); [Greater Accra region] between Weija and Senya Bereku, 20 Mar 1954, J.K. Morton 204 (WAG0185093); [Eastern region] Mamfe, 6 Apr 1955, J.K. Morton 320 (K); [Brong-Ahafo region] Jema, 30 Oct 1955, C.D. Adams 3328 (K); [Greater Accra region] Nungua, 26 Mar 1956, J.O. Ankrah 20166 (K, M, P04577153, W); [Greater Accra region] Nungua, 25 May 1960, R. Rose-Innes 31247 (P04577159); [Greater Accra region] Achimota, May 1961, F.R. Irvine 5474 (K); [Ashanti region] 75 km W of Kumasi, Tano river, 24 Dec 1963, R.A.A. Oldeman 825 (B101143638, BR0000018269888, K, P04577254, WAG0185089); Brong-Ahafo region, between Nkoranza and Kintampo, 27 Dec 1995, C.C.H. Jongkind & C.M.J. Nieuwenhuis 2548 (BR0000018269871, WAG0050916);

Guinea: [Boke region] Monchon, 6 Feb 1979, S. Lisowski 51384 (BR0000018269802);

Guinea-Bissau: Antula, 21 Mar 1943, Anonymous 1499 (B101143640, BR0000018269819, FT0007105, K, M, P04577132, WAG0104159); [Bissagos Islands] Formosa, 26 Apr 1945, Anonymous 1969 (B101143639, K, M);

Ivory Coast: [San Pédro region] San Pédro, 1900, M. Thoiré 123 (P04577229); [Woroba distr.] Mankono, 2 Jul 1909, A. Chevalier 22002 (P04577125); [Bas-Sassandra distr.] 49 km N of Sassandra, nr Dakpadou, 25 Feb 1959, A.J.M. Leeuwenberg 2858 (BR0000018269864, FT0007110, E, K, M, P04577188, U1398743, WAG0176054); [Montagnes distr.] nr Troya, Cavally river, 8 Mar 1962, J.J.F.E. de Wilde & A.J.M. Leeuwenberg 3555 (BR0000018269857, K, WAG0185084); [Comoé distr.] Ayamé, 12 May 1965, L. Aké Assi 8060 (G); [Lacs distr.] Andokoi, 18 Jul 1970, L. Aké Assi 11277 (G); [Lacs dept.] Betrikan stream, 24 Mar 1971, J. Audru 3828 (P04577264); Bas-Sassandra distr., Niénokoué, Feb 1983, N. Stäuble 0861 (G423540); Bas-Sassandra distr., nr Louga, 8 Apr 1973, J. de Koning 1269 (BR0000018269840); [Zanzan distr.] Bouna, 1 May 1989, P. Poilecot 3923 (G);

Kenya: [Lamu county] Witu, Dec 1892, J.W. Gregory s.n. (BM); Sokoke, 14 Apr 1945, AnonymousW. Jeffrey 162 (G); [Mombasa county] Kibarani, 4 Apr 1946, AnonymousM. Jeffrey 515 (K); Kilifi county, 13 Feb 1946, AnonymousW. Jeffrey 466 (G, K); Machakos county, Kiamkere, 25 Nov 1951, Kirrika 154 (B101143637, BR0000018270044, K); [Lamu county] Mukunguyu Lake, 5 Nov 1957, P.J. Greenway & S.P. Rawlins 9449 (FT0007109, K); [Lamu county] Kiunga, 55 m NE Lamu, 6 Aug 1961, J.B. Gillespie 167 (K001394829); Garissa county, 26 km from Garissa on Hagadera Rd, 0°16'S, 39°47'E, ~230 m, 29 May 1977, J.B. Gillett 21195 (K); Rare river, 11 Dec 1979, J.M. Reitsma 413 (BR0000018268782, WAG0318591); Lamu county, Badar Water Pan, 5 Mar 1980, M.Anonymous Gilbert & P. Kuchar 5892 (K); [Coast prov.] Lamu county, Ras Tenewi, 19 Nov 1988, P. Luke & S.A. Robertson 1425 (K); Kilifi county, Mangea Hill, 250 m, 27 Dec 1988, P. Luke 1587 (K); Tana River county, Tana River Primate Reserve, 17 Mar 1990, P. Luke et al. 524 (K); [Kitui county] Mwingi, 23 Jun 2005, P. Kirika et al. 542 (K); Lamu county, Bodhei to Basuba, 26 Jul 2006, L. Festo et al. 2649 (K); [Kilifi county] Malindi, 17 Nov 2010, S.A. Robertson 7871 (K);

Liberia: [Lofa county] Jenneh, 7 Apr 1909, M. Dinklage 2558 (B101143641); [Nimba county] Ganta, 5 Jun 1973, A. Jacques-Georges 27822 (BR0000018269833, M, WAG0185083); Maryland county, Cavally river, 8 Apr 2000, C.C.H. Jongkind & J. Assi-Yapo 4991 (WAG0013107);

Malawi: [Southern region] Chikwawa distr., Lower Mwanza River, 180 m, 3 Oct 1946, L.J. Brass 17931 (K); [Southern region] Chikwawa distr., Lengwe NP, 300 ft, 15 Dec 1970, A.Anonymous Hall-Martin 1168 (K, P04577250); [Central region] Dedza Distr., Chipoka, 15 Mar 1972, A.J. Salubeni 1785 (K); [Southern region] Machinga Distr., Lake Chiuta Harbour, 23 Dec 1984, I.H. Patel & W. Nachamba 1764 (K); [Southern region] Machinga distr., Dinji vill., 8 Jun 1988, A.J.Salubeni & I.H. Patel 5203 (K); [Southern region] Zomba distr., Chilwa Lake, Mchisi island, 7 Nov 1986, A.J. Salubeni & R.B. Kwatha 4814 (K); Southern Region, Mpoto lagoon, Mauzi, Phalombe, 15°39'23"S, 35°49'9"E, 638 m, 7 Dec 2013, H.T. Vhapama et al. 1088 (K);

Mali: Gao Region, Bamba [without date], ex herb. A. Chevalier 42330 (P04577220); [Niger Delta region] Djenné, 29 Jun 1899, A. Chevalier 1193 (P04577226); [Timbuktu Region] Timbuktu, 8 Aug 1927, O. Hagerup 247 (BR0000017461023, K); [Timbuktu region] Saré-Yamou, 8 May 1932, ex herb. A. Chevalier 183 (P04577221); Gao region, Bagoundjé, 5 Jul 1936, M. de Wailly 5079 (P04577225); Gao region, Niger river, 6 Sep 1936, M. de Wailly 5187 (P04577204); nr Bamako, Sotuba, 13 Oct 1989, A. Raynal-Roques 22802 (P04577269);

Mauritania: [Trarza Region] Rosso, 11 Oct 1969, F.N. Hepper 3617 (K, M);

Mozambique: Lorenço Marques [Maputo], Sep 1913, Anonymous Borle 216 (K); Cabo Delgado prov., 12 Sep 1948, Barbosa 2097 (K); [Inhambane prov.] Homoine distr., 8 Sep 1948, M. Myre & M.F. de Carvalho 232 (K); Manica prov., Dombe, 28 Oct 1953, J. Pedro 4493 (K); [Sofala prov.] Beira, 10 Sep 1962, A.R.A. Noel 2487 (K); Sofala prov., Gorongosa NP, 3 Feb 1963, A.R. Torre & J. Paiva 9007 (BR0000017455930, M); Inhaca island, 23 miles E of Lorenzo Marques [Maputo], 200 m, 9 Sep 1964, A.O.D. Mogg 32038 (K); Lorenço Marques [Maputo] prov., Matola, 21 Jul 1965, A. Marques 588 (WAG1103275); Zambezia prov., Shire river, Vila Bocage, 13 Dec 1971, Anonymous Pope & T. Müller 598 (K); Maputo, 10 Oct 1980, P.A. Schäfer 7280 (BR0000017454452, WAG0155677); Zambezia prov., Mamala, 20 Dec 1996, A.R. Torre & M.F. Correia 16648 (BR0000017454469);

Namibia: [Kunene region] Ongonga, 14 Jan 1904, A. Kestilä 30 (H1056354); [Omusati region] Ruacana Falls, 29 Apr 1962, T.T.Kotze 46 (K, M, PRE0823929);

Niger: Niamey, Sep 1957, A. Vaillant 891 (K); 100 km N of Niamey, Niger river, 14 May 1968, C. Geerling & J. Bokdam 2681 (WAG0104161); Niamey, 2 Apr 1987, N. Leman 81 (BR0000018269963);

Nigeria: Lagos, 26 Mar 1896, Miller 72 (K); [Cross River State] Oban, 1912, P.A. Talbot s.n. (BM); [Benue State] Abinsi, 1912, J.M.Dalziel s.n. (BM); [River State] Port Harcourt, Jun 1930, T.D. Maitland s.n. (K); [Edo State] Benin, 1934, W.A. Fairbairn s.n. (BM); Ondo State, Owo, 31 Mar 1943, A.P.D. Jones 3087 (K); [Ogun State] Shasha (Omo) forest reserve, Akila, 30 Jan 1947, C.F. Onochie & V. Emumwen 20687 (K); [Edo State] r[iver] Osse, Iguoriakhi ferry, 29 Jan 1948, J.P.M. Brenan 8925 (K); [Edo State], Okomu Forest Reserve, 27 Feb 1948, J.P.M. Brenan 9169 (K); [Ouo State] nr Ibadan, 12 Mar 1950, R.D. Meikle 1261 (K, P04577150); [Lagos State] Ikeja, 30 Dec 1952, C.F.A. Onochie 26671 (K); [Oyo State] nr Eruwa, 22 Apr 1958, D.J. Hamber 430 (K, P04577151); [Cross River State] Ebom, 28 Jun 1955, R.H. Stone 26 (K); [Oyo State] Gambari Forest Res., SW of Ibadan, 16 Jan 1958, de Wit 770 (WAG0185105); [Kogi State] Lokoja, 5 Jun 1958, B.O. Daramola 36930 (K, WAG0185106); [Niger State] Shagunu, 10 km N of Bussa, 31 Jul 1965, C.D.K. Cook 471 (K, P05307013); [Kogi State], Koton Karfe, 27 Feb 1968, B.O. Daramola & A. Binuyo 61909 (K); Cross River State, Afunatam, 30 Mar 1972, J. Lowe 13580 (K); [Cross River State] Ikom, Agbokim waterfalls, 26 Feb 1973, Latilo & Oguntaya 67685 (K. WAG0185111); [Ekiti State], Igbara Odo, Oruwo stream, 9 Mar 1973, Olorunfemi & Fagbemi 70748 (K, WAG0185110); [Niger State] Minna, Gurara Waterfalls, 17 May 1973, Eimunjeze et al. 66411 (WAG0185108); Oyo State, Ibadan, Asejire, 4 May 1974, Z.O. Gbile 73431 (K, WAG0185112); [Edo State], Iguoriakhi, 16 May 1974, Eimunjeze & Oguntayo 70213 (K);

Republic of Congo: Niari dept., 29 Nov 1951, J. Koechlin 1587 (P04576681); Ubangi river, 20 Feb 1963, de Nere 907 (P04577259); Moutou ya N’Gombé, 12 May 1968, P. Sita 2036 (P04577256); [Pool Malebo] Mbamu, 14 Oct 1969, F. Hallé 1641 (P04577257); nr Brazzaville, 26 Aug 1969, Y. Attims 205 (WAG0034944); nr Brazzaville, 15 Apr 1970, Y. Attims 442 (BR0000018270020); Kouilou Dept., 20 km SE of Pointe-Noire, Cayo Lake, 22 Mar 2017, E. Bidault & al. 3022 (BRLU0018798);

Senegal (selected): “Waalo”, 1831, Perrottet s.n. (G); Dakar, 9 Apr 1948, J.Anonymous Adam 1053 (WAG1103302); [Thiès region]Thiès, May 1948, R.P. Berhaut 924 (P04577233); [Dakar region] Mbao, 1950–1951, R.P. Berhaut 2020 (BR0000018269703, P04577235); [Dakar region] Dagoudane-Pikine, 23 Apr 1960, J. Raynal & A. Raynal 5726 (P04577242); [Thiès region] Kayer, 16 Jun 1961, J. Raynal & A. Raynal 7067 (P004577267); [Ziguinchor region] Bignona, 31 Mar 1964, R.P. Berhaut 7267 (BR7000577);

Sierra-Leone: [Northern prov.] nr Madina, 11 Apr 1892, AnonymousF. Scott Elliot 5561 (BM, K); [Northern prov.] Mokele, Sep 1914, Anonymous s.n. (BM); [Southern prov.] Juring, 1 Dec 1926, F.C. Deighton 297 (K); [Northern prov.] Makump, 3 May 1929, F.C. Deighton 1707 (BM, K); [Southern prov.] Baoma, 16 Apr 1936, F.C. Deighton 3166 (K); [Northern prov.] nr Kasanko, 13 May 1951, P. Adams 225 (K); [Southern prov.] Ngokuma (Kori), 22 Jun 1952, F.C. Deighton 5845 (K); [Northern prov.] nr Kambia, Magbema, 25 May 1954, H.D. Jordan 958 (K); [Western Area prov.] Fogbo, 3 Mar 1964, Morton & Jarr 917 (WAG0185090);

Somalia: [Puntland State] Ganaane to Marro Mogale Umberto I, Mar 1893, D. Riva 731 (FT0007112) [Note. This specimen was collected in the same locality as FT0007113 (var. lanatus)];

South Africa: [KwaZulu-Natal prov.] [Port of] Natal [Durban], 1847, Guienzius 138 (S, G, sub Mollugo serrulata Sond.); [KwaZulu-Natal prov.] Isipingo, 22 Jun 1893, B. Schlechter 2804 (BOL217408, FR-0132687, G, P04577164); Durban, 19 Oct 1894, M. Wood s.n. (P04577180); [KwaZulu-Natal prov.] Maputaland, Jun 1914, Anonymous 14294 (BOL217409); [KwaZulu-Natal prov.] Umbilo, nr Durban, 4 Dec 1919, C.E.Moss 6657 (K); [KwaZulu-Natal prov.] Port Shepstone, 9 Aug 1919, R.G.Strey 6444 (BR0000017454520, K, M); [KwaZulu-Natal prov.] Durban, 20 Sep 1965, C.J. Ward 5148 (K); [KwaZulu-Natal prov.] Ndumo Game Reserve, 9 Jan 1969, E.S. Pooley 364 (E); Kruger NP, [Mpumalanga prov.] Crocodile river, nr Komatipoort, 7 Apr 1992, van Rooyen & Bredenkamp 518 (PRE0784699); [Mpumalanga prov.] Kruger NP, Skukuza, 5 Jan 2006, F. Siebert & S.J. Siebert 146 (PRE0851635);

South Sudan: [Sudd region] Bahr el Zeraf, 26 Apr 1930, N.D.Simpson 7790 (BM, K); [Upper Nile State] [Upper Nile State] Sobat river, nr Nyanding, 7 Jun 1929, D. Simpson 7085 (BM, K); [Upper Nile State] Shambe, 14 Jul 1929, D. Simpson 7380 (BM, K);

Swaziland: Blue Jay Ranch, Lubombo Mountains, 30 Oct 1977, J.Culverwell 1114 (K, PRE0498323);

Tanzania: Zanzibar, 1847, L.H. Boivin s.n. (P04577175); Zanzibar, Jul 1873, J.M. Hildebrandt 907 (BM, LE); Kilimanjaro region, [Moshi Rural distr.] Kahe, Apr 1895, G. Volkens 2188 (E, LE); Pemba [island], [without date] J.H. Vaughan 899 (K); [Lindi region] Masoko, 4 Nov 1912, A. Stolz 1649 (K); [Mbeya region] Kyimbila, N of Lake Nyasa, 1915, A. Stolz K13 (BM, BOL217411, BR0000017454384, K); Tanganyika terr., [Pwani region] Rufiji, 3 Jan 1931, Anonymous 168 (K); Tanga region, Amboni, 6 Jul 1932, Geilinger 661 (K); [Lindi region] 40 km W of Lindi, Lutamba Lake, 29 Nov 1934, H.J. Schlieben 5660 (B101143615, BM, BR0000017454377, G); [Katavi region] Lake Katavi, 3500 ft, 25 Jan 1950, A.A. Bullock 2334 (K); [Mwanza region] 3700 ft, 24 Oct 1952, R.E.S. Tanner 1100 (BR0000017454322, FT0007111, E, K, WAG1103308); [Tanga region] Mkomazi river, 2 miles NE of Lake Manka, 440 m, 1 May 1953, R.B. Drummond & J.H. Hemsley 2347 (B101143612, BR0000017454391, FT0007106, K); Tanga region, Pangani, 500 ft, 23 Jul 1955, R. Tanner 1982 (K); Tanga region, Pangani, 27 Nov 1955, R. Tanner 2369 (BR0000017454339, K); [Pwani region] Utete by river Rufiji, 120 m, 2 Dec 1955, E. Milne-Redhead & P. Taylor 7527 (B101143610, BR0000017454360, K); [Morogoro region] Ilonga, Kilosa, 1700 ft, 18 Jan 1958, M. Cole 58 (K); [Kigoma region] Lake Tanganyika, Kibwesa point, 6°30'S, 29°57'E, 3000 ft, 20 Aug 1958, T.G. Jefford & J. Newbould 1673 (K); [Kigoma region] Kungwe Mt, Kasoje, 2600 ft, 17 Jul 1959, J. Newbould R.M. Harley 4408 (B101143609, BR0000017454315, K); [Tanga region] Nzega distr., Wembere escarpment, 1200 m, 31 Oct 1960, H.M. Richards 13452 (K); [Morogoro region] Kilosa distr., Mazunyungu river, 25 Nov 1961, S.R. Samsei 3443 (K); Dodoma region, 1 mile S of Dodoma, Imagi hill, 4000 ft, 29 Jan 1962, R. Polhill & S. Paulo 1290 (B101143611, BR0000017454292, K, P04577203); [Dodoma region] Mtera, 19 Apr 1962, R. Polhill & S. Paulo 2077 (B101143604, BR0000018268904, K); Zanzibar, Migombani, 18 Aug 1962, Faulkner 3090 (BR0000018269062, K); [Tanga region] Lushoto, Korogwe stream, 15 Dec 1962, M.E. Archbold 83 (K); Arusha region, Ngurdoto NP, 1650 m, 13 Apr 1965, M. Richards 20178 (BR0000018270082, K); [Arusha region] Longil Lake, Ngurdoto Crater NP, 4900 ft, 24 Feb 1966, P.J. Greenway & Kanuri 12385 (BR0000017454278, K); Arusha NP, Lake Kawanya ya Matteo, 1524 m, 4 Nov 1969, H.M. Richards 24592 (BR0000018270089, K); Arusha region, Lekuruki vill., 914 m, 5 Dec 1969, H.M. Richards s.n. (BR0000018270136, K); [Morogoro region] Kilosa, at Ruaha river within 1 km E of Kidatu bridge, 400 m, 17 Jul 1970, M. Thulin & B. Mhoro 478 (K); [Katavi region] Kapapa, 974 m, 19 Sep 1970, H.M. Richards & S. Arasululu 25994 (BR0000018270112, K); Arusha NP, Ngongongare Maji-ya-Punda gorge, 4900 ft, 1 Feb 1971, H.M. Richards & S. Arasululu 26625 (BR0000018270129, K, M); [Morogoro region] Kidatu, 7°7'S, 36°9'E, 14 Jan 1971, B. Mhoro 178 (BR0000018270075, K); Arusha distr., Lekuruki area, 27 Mar 1971, H.M. Richards & S. Arasululu 26866 (BR0000018270105, K); Iringa region, Ruaha NP, 790 m, 26 Jan 1972, A. Bjornstad 1274 (K); [Lindi region] nr Kingupira, 25 Jun 1975, K. Vollesen 2490 (WAG1103312); Tanga [region] Tanga, 1 Oct 1979, F.C. Magogo 2000 (BR0000017454261); Tanga region, 4 Sep 1987, M.E. Archbold 8145 (K); [Tanga region] Kwamkono, 20 Sep 1990, M.E. Archbold 3309 (K); Kilimanjaro, 7 Apr 1994, J.M. Grimshaw 94334 (K); [Tanga Region] Kwasunga, 23 Mar 2012, M. Alvarez 921 (B100599491);

Togo: Lomé, [years] 1900–1902, Warnecke 299 (BM, BR0000018269895, G, K, P04577158); Togo Lake, 29 Jan 1978, H. Scholz 2955 (B101143632, K, P04577245); Lomé, 15 Apr 1978, M. Hakki et al. 48 (B101143635, K); [Kara region] NW of Bassar, 25 Apr 1978, M. Hakki et al. 507 (B101143634, K, P04577246); Lomé, 17 Feb 1979, A.J.M. Leeuwenberg 11935 (BR0000018269918, WAG0185086); Aneho, 6°14'N, 1°37'E, alt. 30 m, 26 Jun 1985, Pasch 8626 (K); [Maritime region] Aného, 26 Jun 1985, P. Schäfer 8626 (B101143633, WAG0185087);

Uganda: Victoria Lake, Lolui Island, 11 May 1965, G. Jackson 511565 (K); [Western region] Masindi distr., Bunyoro, 1050 m, 15 Feb 1999, K.A. Lye & A.B. Katende 23474 (K);

Zambia: [Western prov.] Kande Lake, 8 miles NE of Mongu, 11 Nov 1959, R.B. Drummond & A.J.Cookson 6335 (E, K); [Muchinga prov.] [Northern prov.] Mweru-wa-Ntipa, Katema island, 8°50'S, 29°40'E, 2950 ft, 25 Jul 1962, P.J.Tyrer 109 (BM, BR0000017454421); [Northern prov.] Abercorn [Mbala] Distr., Sumbu Bay Lake, 780 m, 29 Dec 1963, M. Richards 18710 (K); [Lusaka prov.] [Muchinga prov.] Mpika Distr., Mfuwe Camp, 2000 ft, 12 Dec 1968, W.L.Astle 5389 (K); [Northern prov.] Kaputa distr., Mweru Wantipa NP, 1000 m, 8 Dec 1993, M. Marello et al. 989 (WAG0023616);

Zimbabwe: [Masvingo prov.] Chivi, Lundi river, 2500 ft, 30 Oct 1955, H. Wild 4692 (BR0000017454445, K); [Manicaland prov.] Chipinge, S of Giriwayo camp, 1230 ft, 13 Mar 1957, J.B. Phipps 616 (K).

Tropics of the Old World. Additionally, we report here the first record from North Africa [Egypt: [Sinai Peninsula] Wadi Isleh, Jan 1903, R. Muschler s.n. (G!)].

Dry river beds and other water bodies, or as a weed.

A variety with ± villose pubescent stems, shorter leaves (up to 10 mm) and shorter (up to 5 mm) pedicels (Fig. 25). Some individuals also have shorter perianths (2.7–3.6 mm) and anthers (0.3–0.6 mm) as well as brighter (yellow-brown) seeds compared to the type variety, but these characters are not always diagnostic.

This interpretation and circumscription of the variety is undertaken for the first time. In Africa, plants known as G. microphyllus Hauman were considered endemic to DR Congo (Hauman 1949; Lebrun and Stork 2003). Phuphathanaphong (2005) reported G. oppositifolius var. keenanii (as G. herniarioides) from Thailand, but described it as stellate-pubescent. However, the type specimens have simple hairs only, and it is not clear which name should be applied to the Asian plant reported as “G. herniarioides”.

Gagnepain (1918) cited two specimens from Vietnam collected by B. Balansa (no. 3397 & 4613). We found only duplicates numbered 4613 (Environs de Hanoi, dans les rizières après la moisson, June 1891, B. Balansa 4613), which were distributed from Paris to various herbaria (K000641793!, K! [without a barcode], E!, NY00288837, US00103110), whereas another gathering (Balansa 3397) was traced at P and is designated as lectotype here.

Ewart and Petrie (1926) reported that both Wycliffea obovata (=Glinus oppositifolius s.str.) and W. rotundifolia (= G. oppositifolius var. keenanii) were described from the same locality in Australia. The specimens collected by D. Riva in Somalia [Ganaane to Marro Mogale Umberto I, Mar 1893] also belong to different varieties: the specimen FT0007113 is a typical var. keenanii, while FT0007112 represents var. oppositifolius. Such plants with shorter leaves and pedicels are found in various localities across the distribution area of G. oppositifolius: in Australia, India, Vietnam, many African countries (Gilbert 2000; Fig. 26 in the present article). A specimen of G. oppositifolius var. keenanii used in the molecular study falls within G. oppositifolius subclade (Figs 1, 2). We prefer to consider such plants as a variety of G. oppositifolius, perhaps growing under less favorable drier conditions.

(Fig. 26). Angola: Namibe prov., Tombe [Tombua], 15 Jan 1956, E.J. Mendes 1301 (LISC032017);

Benin: [Alibori dept.] Malanville, 26 Apr 1999, P. Houngnon et al. 6501 (BR0000017456159, WAG0235844);

Cameroon: North prov., Bénoué, 22 May 1974, C. Geerling & J. Néné 4858 (BR0000017456128, WAG0330419);

DR Congo: see the type specimens of G. microphyllus and G. oppositifolius var. lanatus;

Ethiopia: [Oromia region] Illubabor zone], S of Abobo, 650 m, 14 Apr 1982, I. Friis et al. 2456 (K);

Kenya: Kwale county, 31 Aug 1953, R.B. Drummond & J.H. Hemsley 4068 (BR0000018268805, FT0007108, K);

Malawi: [Southern Region] Shire Highlands, Jul 1885, J. Buchanan s.n. (E, K); [Southern region] Njobru, Mulanje, 17 Nov 1955, G. Jackson 1765 (BR0000018269130, K);

Mozambique: Gaza prov., Baxo Limpopo, 27 Aug 1969, M.F. Correia & A. Marques 1280 (WAG1103322); [Sofala prov.] Beira, Jul 1970, K.L. Tinley 1936 (K, M); Inhambane prov., Vilankulos, 18 Sep 1973, M.F. Correia & A. Marques 9417 (WAG1103327); Cabo Delgado [prov.], Ruvuma riverbank, 10°34'39"S, 40°21'11"E, 10 m, 15 Nov 2009, Q. Luke 13803 (K);

Somalia: [Puntland State] Ganaane to Marro Mogale Umberto I, Mar 1893, D. Riva 721 (FT0007113).

Sudan: [State of Sennar] Sennar, May 1867, A. Figari s.n. (FI059439);

Tanzania (new records): Morogoro Region, Magogoni, Aug 1930, A. Haarer 1839 (K); Rukwa Region, Rukwa, 2800 ft, 5 Dec 1954, H.M. Richards 3530 (K); [Tanga Region] Handeni distr., Horogwe to Handeni, 18 Nov 1955, E. Milne-Redhead & P. Taylor 7337 (BR0000018269048, K); Rufiji distr., 2 Dec 1955, E. Milne-Redhead & P. Taylor 7526 & 7528 (BR0000018269031, BR0000018269055, K); [Manyara region] Mbulu distr., Tarangire NP, 1066 m, 24 Nov 1969, M. Richards 24759 (BR000017456005; K);

Zambia: [Eastern prov.] Petauke distr., Luangwa river bridge, 5 Sep 1947, P.J.Greenway & J.P.M.Brenan 8045 (K); Chinsali distr., Mbesuma, 11 Oct 1960, E.A. Robinson 3950 (K, M); Katondwe, 14 Nov 1963, D.B. Fanshawe 8129 (K); Luangwa valley, 4 Dec 1965, W.L. Astle 4166 (K); Western Province, Bulozi plain near Mongu, 15°10'S, 23°06'E, 1000 m, Oct 1993, M.G. Bingham 9743 (K, WAG0334359);

Zimbabwe: [Masvingo prov.] Nuanetsi, nr Malipati, 2 May 1961, R.B. Drummond & R.O.B. Rutherford-Smith 7677 (K);

Tropical Africa (incl. Madagascar: WAG!), Asia, Australia.

Angola. Pungo Andongo distr., [by a nearly dried swamp between Condo and Quisonde] 2400–3800 ft, [end of March 1857] F. Welwitsch 1111 (LISU, n.v.; isotypes BM!, BR!, K000221394!, K000221395!).

The type label has been complemented with the information from Hiern (1898: 416).

Originally (Gonçalves 1965), the name G. oppositifolius var. glomeratus was invalidly published with a diagnosis in Latin based on two gatherings without explicit type designation. It was validated later (Gonçalves 1970) by indicating a “lectotype” and providing a full and direct reference to the validating diagnosis.

This variety is characterized by the clusters with >10 flowers, and sessile or shortly (up to 4.0 mm) pedicellate flowers (Fig. 27). It may represent a montane variety of G. oppositifolius growing at higer altitudes (1000–2000 m), because one of the specimens cited by Gonçalves (1965) from Bié province, Angola, was collected at the elevation of 1360 m a.s.l. However, we did not include the samples of this variety into the molecular analysis, and further studies are needed to prove its varietal rank.

The plants of G. oppositifolius var. glomeratus look similar to G. hirtus. However, at least two characters distinguish it from the latter species: simple curved hairs vs. stellate pubescence, and shorter (0.3–0.6 mm long) stylodia (as in the type variety) vs. 0.5–1.0 mm long stylodia in G. hirtus. This variety was reported from Angola (Gonçalves 1970), but here we extend its distribution into DR Congo, Tanzania and Zambia.

(Fig. 26). Angola: [Cuanza Sul prov.] between Condo and Quisonde [Porto Amboim], Mar 1854, F. Welwitsch 1111 (BM); Cuando-Cubango prov., Longa, Longa river, 1360 m, 17 Mar 1960, E.J. Mendes 3155 (BM013839321, LISC032016, M);

DR Congo (new records): [Kongo Central prov.] Kisantu, Kingimba Kinga, 3 Oct 1958, L. Paulwels 108 (BR0000017454933); [Haut-Katanga prov.] 26 km NW of Lubumbashi, Lupoto, 2 Mar 1961, A. Schmitz 7129 (P05307011); Haut-Katanga prov., Kundelungu Plateau, 1700 m, 24–29 Mar 1969, S. Lisowski et al. 4053 (BR0000018268348);

Tanzania (new records): Shinyanga Region, Shinyanga, 1932, R.D. Bax 23 (K);

Zambia (new records): [Luapula prov.] Kawambwa, 23 Aug 1957, D.B. Fanshawe 3561 (K); [North-Western Province] Mwinilunga District, Mujileshi river, 1290 m, 8 Nov 1962, H.M. Richards 16953 (K).

Tropical Africa (south of the equator).

Jeffrey (1961: 16) indicated the “type” of Glinus setiflorus at C. He had not seen the material, and failed to distinguish between several sheets in Forsskål’s collection. We designate one of these specimens as lectotype, which was seen and annotated by P.Ascherson in 1881.

The locality information and collection date are complemented from Hepper and Friis (1994).

(Fig. 28). Annual with prostrate or ascending stems densely covered with persistent stellate hairs; prickles absent. Leaves shortly petiolate (petioles 2–15 mm long), grey or white, farinose, entire or slightly crisp, 10.0–35.0 × 7.0–25.0 mm, obovate to almost roundish, veins adaxially recessed and abaxially prominent. Flowers in clusters of (1) 2–4, sessile or with pedicels up to 5(10–15) mm long. Perianth segments 5, ovate, in flowering 6.0–7.0 mm long, in fruiting 9.0–11.0 × 7.0–8.0 mm, with a mucro 0.5–1.0 mm long; white or creamy inside, abaxially with dense stellate hairs; flower buds and anthocarp ovoid or almost roundish (Fig. 5A); yellowish multifid petaloids often present. Stamens 5; anthers 0.8–1.0 mm long. Stigmas 5, 0.7–1.2 mm (sometimes a style is present up to 0.7 mm long). Seeds 0.7–0.9 × 0.50–0.60 mm, almost black, colliculate and without longitudinal striae; aril hood often reduced, rarely clearly visible, and in the latter case up to 0.30 mm long.

Sands, seasonally flooded depressions, rocks, limestones or clayey soils; 0–1300 m.

(Fig. 29).

Ethiopia: [Oromia Region] Adama, 23 Dec 1913, D. Riva 1999 (FT0007119); [Somali region] Ogaden distr., Danot, 2 Dec 1944, P.E. Glover & H.B. Gilliland 440 (BM, K); [Afar Region] Afambo, May 1953, E. Chedeville 13 (FT0007028 – right specimen); [Somali region] Ogaden, nr Mardere, 2 Dec 1953, G. Popov 1143 (K); [Somali region] Balbeh Danot, 30 Jun 1958, C.F.Hemming 1486 (FT0007118, K); [Somali region] N of Gara Afdem Mtn., 9°38'N, 40°50'E, 20 Jan 1964, W. Burger 3417 (FT0007116, K, WAG0185097);

Kenya: Ewaso Ng’iro river, 2 May 1945, P.R.O. Bally 4506 (K); [Isiolo county] Kipsing, 2 May 1945, P.R.O. Bally 4506 (G); [Mandera county] Banissa, 1020 m, 22 May 1952, J.B. Gillett 13257 (BR0000018268812, FT0007120, K); [Isiolo county] Tana River distr., nr Garba Tula, 18 Jul 1952, P.R.O. Bally 8224 (FT0007125, K); Garissa county, S of Garissa, 26 Sep 1957, P.J. Greenway 9236 (FT0007121, K); [Mombasa county] Mombasa, 30 Oct 1959, D. Napper 1343 (BR0000017456050, K); [Tana river county] Kamor Jila, 1°21'S, 39°55'E, 12 Dec 1964, J.B. Gillett 16414 (BR0000017456067, FT0007123, K, P04577112, WAG1103335); [Taita-Taveta county] Tsavo NP, 2 Jan 1967, P.J. Greenway & Kanuri 12937 (FT0007122, K); Turkana county, Lomesekin, locally very common, 29 Aug 1968, O.M. Mwangangi & D. Gwynne 1218 (K); Turkana county, Lodete, 570 m, 24 Jul 1969, O.M. Mwangangi 1378 (FT0007124, K); [Mandera county] 54 km NE of El Wak, 380 m, 12 Dec 1971, Bally & Smith 14572 (K); Samburu county, nr Wamba, 3 Jul 1974, R.B. Faden & A.J. Faden 74-946 (BR0000017456043, K, WAG1103337); Garissa county, Garissa to Hagadera, 230 m, 29 May 1977, J.B. Gillett 21193 (FT0007126, K); [Tana River county] Kora camp junction, 3 Sep 1983, G.M. Mungai 83/35 (K); [Makueni county] Chyulu plains, Kigelia, 26 Jul 1992, Luke 3204 (K); [Kitui county] Ukasi Dam, 3 Aug 2006, L. Festo & Q. Luke 2813 (K, LE); Makueni county, Kibwezi, 990 m, 22 Nov 2010, Q. Luke 14955 (K);

Somalia: [Sanaag region] nr Dofar [El Dahir], 19 Oct 1954, P.R.O. Balley 10115 (FT0007117, K); [Lower Juba Region] Lag Badana, 28 Oct 1971, G. Moggi & R. Bavazzano s.n. (FT0007034); [Bay region] Dinsoor, 16 Feb 1981, J.J. Beckett 1491 (K); [Galguduud region] 27 km S of Guri Ceel, 5°05'N, 45°47'E, 6 Jun 1988, P. Kuchar 17836 (K);

Sudan: Kassala State, S of Tob el Ahmar, 16 Apr 1961, J.K. Jackson 4143 (K);

Tanzania: [Dodoma region] 29 miles S of Dodoma, 3500 ft, 15 Aug 1928, P.G. Greenway 773 (BM, K); Dodoma region, Ikowa Dam, 60 km E of Dodoma, 900 m, 29 Jul 1970, M. Thulin & B. Mhoro 515 (K001394832); Dodoma county, Ikowa reservoir, 980 m, 19 Jun 1974, B. Mhoro & I. Backéus 1949 (WAG0192068).

East Africa; Arabian Peninsula (Yemen). In addition to the records cited, it is apparently present in Djibouti and Eritrea. In Yemen, where G. setiflorus was described from, it is only known from the type locality (Miller 1996).

Holotype: Zambia. Barotseland, [Western province] Mongu, mud at edge of river, 6 Jan 1966, Robinson 6780 (K000865904! iso – M!).

Annual up to 70 cm, often forming mats, with numerous prostrate or ascending stems, covered at least in young parts with dense stellate hairs, so that the plant appears greyish; prickles absent. Leaves in false whorls of 2–5, oblong-spatulate or obovate, rarely almost roundish, entire, 10.0–40.0(50.0) × 5.0–15.0(20.0) mm, shortly pedunculate (peduncles up to 15.0 mm), green, scarcely pubescent, rarely subvillose, veins not prominent abaxially. Flowers in clusters of 2–12, pedicellate (pedicels 5.0–20.0 mm long, rarely shorter). Perianth cylindrical (Fig. 5B), perianth segments (5) oblong or ovate, 5.5–7.0 mm long at anthesis, 2.5–3.0 mm wide, abaxially with scattered stellate, easily caducous hairs (at least along mid-vein), abaxially green or pink at senescence, adaxially white or creamy, in fruiting reaching 6.5–8.0 mm long, with a mucro 0.5–1.0 mm long. Stamens 10–15, outer stamen series often sterile with filaments terminating with teeth; anthers (1.0)1.2–1.8 mm long (1.0–1.2 mm in plants from higher altitudes). Stylodia 3, 1.0–1.5 mm long (sometimes united in the lower half into a style). Seeds reddish or brown-red, 0.5–0.6 × 0.35–0.50 mm, colliculate, without longitudinal ridges, aril hood clearly visible, ca. 0.25 mm long.

In the herbaria, G. zambesiacus was identified as G. lotoides, G. lotoides var. virens, G. lotoides × G. oppositifolius, and sometimes as G. bainesii. Glinus zambesiacus, G. oppositifolius, G. bainesii and G. hirtus are found in similar habitats. All character sets for each species are indicated in Table 5. The new species can be easily differentiated from both G. oppositifolius and G. bainesii by indumentum consisting of stellate trichomes, while G. hirtus has much shorter perianth and fewer (up to 8) stamens.

Glinus zambesiacus is related to African G. lotoides and G. setiflorus (Figs 1, 2) and is easily differentiated from both by cylindrical anthocarp, not recessed leaf veins, length of anthers, number of stigmas, and seed colour (see Table 5).

Sands, savannas or as a weed at altitudes up to 2200 m a.s.l. The plants growing in montane areas (DR Congo: Katanga province; Kenya; Tanzania) have much shorter pedicels (up to 6.0 mm long), and their flowers are subsessile. Such specimens were labelled in some herbaria as Glinus zambesiacus var. brevipedicellatus Sukhor. but this character seems to be variable in the specimens collected at lower altitudes.

The new species is named after the territory of Zambezi river basin as its main distribution area.

(Fig. 31). Angola: [Cunene prov.] Mucope, 28 Nov 1963, C. Henriques 261 (BM, K, LISC032006);

Botswana: Ngamiland [North-West distr.], E of Cuando river, 3100 ft, Oct 1945, H.H. Curson 1052 (PRE0403680);

DR Congo: [Katanga prov.] Pweto, 952 m, 17 Apr 1926, W. Robyns 1999 (BR0000018269352, WAG0185132); Katanga prov., Mitwaba territory, Upemba, Sep 1948, G.F. de Witte 4304 (K, M, P04576569, WAG0185135 Katanga prov., Kasenga, 1 Nov 1948, A. Schmitz 2101 (BR0000018269376); Katanga prov., Kasenga, 920 m, 2 Jan 1960, J.-J. Symoens 7048 (BR0000018269680); Katanga prov., Pweto, 900 m, 18 May 1971, F. Malaisse 6767 (BR0000018269567); Katanga prov., Kantala river, 1480 m, Apr 1975, F. Malaisse 8724 (BR0000018269345, WAG0185133);

Kenya: [Nakuru county] Naivasha, 6000 ft, Aug 1934, A. Turner 6646 (K); [Nakuru county] Naivasha Lake, 6450 ft, Apr 1938, Team of Dept. Agric. Uganda 2351 (B101143600, BR0000018268751, K); nr Nairobi, 6000 ft, May 1943, G. Babault 5089 (FT0007102, K); [Trans-Nzoia county] Karara Dam, 13 Apr 1958, Y.E. Symes 339 (K); [Kajado county] Suswa, 6000 ft, 27 May 1963, P.E. Glover 3717 (BR0000017455992, K); [Nakuru county] Lake Naivasha, 22 Oct 1976, T. Hayes 160 (K); [Nakuru county] NW of Lake Naivasha, 1890 m, 15 May 1982, O.M. Mwangangi 2282 (K);

Namibia: [Oshana region] Amboland, Ongonga [Ondangwa], 8 Feb 1907, E. Liljeblad 80 (H1003175); [Kavango West region] Junction of Mpungu Omuramba and Okavango River between Tondoro and Lupala, 17 Dec 1955, B. de Winter 3974 (K, M); Namibia, Caprivi [Strip], Chobe river, 12 Nov 1970, G. Mahundu 431 (PRE0736547);

Zambia: [Western prov.] N of Kalabo, Barotse, Omboya stream, 11 Nov [without year], H.J.A. Rea 146 (K); [Central prov.] Luombwa, 16 Nov 1930, G.F. de Witte 37 (BR0000018269321, BR0000018269338); [Luapula prov.] Bangweulu Lake, May 1946, Bredo 1980 (BR0000018269529); [Eastern prov.] Petauke distr., Lunagwa river, 5 Sep 1947, P.J. Greenway & J.P.M. Brenan 8052 (K); [Western prov.] Shangombo, 3400 ft, 9 Aug 1952, L.E. Codd 7463 (BM, K); [Copperbelt prov.] Kitwe, 27 Dec 1955, D.B. Fanshawe 2679 (BR0000018269574, K); [Southern prov.] Namwala, on Kalahari sand, waste ground nr gardens, 9 Jan 1957, E.A. Robinson 2101 (BR0000018269581, K); [Luapula prov.] Kawambwa, 10 Nov 1957, D.B. Fanshawe 3894 (K); Mongu, 11 Nov 1959, R.B. Drummond & A.J. Cookson 6334 (E, K); [Luapula prov.] Fort Rosebery [Mansa], 8 Feb 1959, R. Watmough 231 (BM); Western Province, Bulozi plain, 22 Dec 1959, W. Gilges 748 (K); [Copperbelt prov.] Kaluba, 940 m, 24 Dec 1959, J.-J. Symoens 6822 (BR0000018269598); [Muchinga prov.] Mbesuma, 11 Oct 1960, E.A. Robinson 3950 (M); [Central prov.] Mwenda, 12 Dec 1960, J.-J. Symoens 7942 (BR0000018269383); [Northern prov.] Kasama, Chambesi river, 1200 m, 7 Apr 1961, H.M. Richards 15023 (K); [Northern prov.] Mofwe Dambo, 2850 ft, 30 Jul 1962, P.J. Tyler 148 (BM); [Lusaka prov.] Kafue, Kitwe, 11 Dec 1962, J.M. Mutimushi 237 (K); [Southern prov.] Namwala, 17 Dec 1962, H.J. van Rensburg 1105 (K); [Luapula prov.] Ndoba to Lubwe, Fourré station, 1140 m, 27 Jan1963, J.-J. Symoens 10023 (BR0000018269123, K); [Luapula prov.] W of Lake Bangweolo, 2 km N of Mwamfuli, 23 Dec 1964, J.J. Symoens 11202 (K); [Eastern province] Luangwa valley, Mfuwe, 26 Feb 1965, W.L. Astle 4121 (K); [Western province] Bulozi [Barotse] Plain nr Mongu, 14 Oct 1993, M.G. Bingham 9740 (K); [Luapula prov.] Samfya, 11°21'S, 29°33'E, 21 Apr 1989, G. Pope et al. 2194 (BR0000018269093, K); [Lusaka prov.] Dembo Chisangwa, nr Kafwe [Kafue], 1140 m, 23 Nov 1998, B. Leteinturier et al. 439 (BR0000018269079 & BR0000018269086);

Tanzania: [Mara region] Musoma distr., E of Naabi Hill, 10 May 1961, P.J. Greenway & al. 10160 (K); [Ndutu region, nr Naabi Hills] Lamuta Hills, 5000 ft, 22 Jul 1962, J. Oteke 201 (K); [Arusha region] Ngorongoro Crater, 17 Jan 1965, J. Goddard 196 (WAG1103310);

Zimbabwe: [Matabeleland North prov.] Victoria Falls, Feb 1906, C.E.F. Allen 284 (K); Victoria Falls, 5 Jan 1955, E.A.Robinson 1058 (K); [Matabeleland North prov.] Sebungwe, Zambesi river, Binga, 1460 ft, common, 9 Nov 1958, J.B. Phipps 1414 (BR0000018269154, K).

The images of B.T. Wursten taken in Mana Pools, Zimbabwe, and Kandahar Fishing site [nr Livingstone, Zambia] for the online “Flora Zambesiaca” (as G. lotoides var. virens) are G. zambesiacus.

The species is only known from the countries listed above.

The American Glinus radiatus was reported for Senegal and some other African countries (Central African Republic, Mali, Nigeria, and South Africa) as an alien species originating from tropical America (Berhaut 1979). It was later reported for Mali, Niger, Central African Republic, and repeatedly for Senegal by Boudet et al. (1986), for Benin by Akoegninou et al. (2006), for Guinea, Sierra Leone, Ghana, Ivory Coast by Lisowski (2009) and for Burkina Faso by Thiombiano et al. (2012) and Schmidt et al. (2017). We did not see any specimens of true G. radiatus from the whole of Africa, and almost all specimens identified as G. radiatus belong to the morphologically similar G. hirtus. The differences between them are mentioned in the notes under G. hirtus. One of the specimens from Burkina Faso (Gnagna prov., Bogande, FR-0019665) identified as G. radiatus is indeed Zaleya pentandra (L.) C.Jeffrey (Aizoaceae).

The protologue (Forsskål 1775) was based on the only specimen indicated by Hepper and Friis (1994) as holotype. Hartmann (2001: 31) designated figure 14 in Forsskål (1776) as lectotype but this choice has no standing.

The locality information and the collection date are complemented from Hepper and Friis (1994).

n.v. Probably described from Mozambique.

No specimens were cited in the protologue (Sprengel 1825), and the description of G. mozambicensis is short and cannot be evaluated properly. According to Don (1834) and Endlicher and Fenzl (1839), it is a synonym of Gisekia pharnaceoides L. sensu lato [incl. G. africana Kuntze] (Gisekiaceae).

Neotype (designated here): Hb. Wight in Wallich Cat. 1543 (addenda) (K001113401!).

Roth (1821) is an account of plants collected by Benjamin Heyne in India. The corresponding herbarium specimens were donated to Roth, whose collection was purchased by the Berlin Botanical Garden (B) and largely destroyed together with the holdings of that Herbarium in 1943 (Hiepko 1987). Heyne made no records on the exact provenance of his specimens, and “India Orientalis” (the Indian Peninsula) is the only information published and known.

In the absence of any surviving original material, we used for neotypification one of the classical specimens assigned to the species by Fenzl (1836), who was the first to interpret the name.

Heyne (in Roth 1821) described G. trianthemoides as a plant with glabrous stems, obovate leaves with rounded tips, and lax, almost dichotomous inflorescences. Such characters are not found in any Glinus. Wight (Wight and Walker-Arnott 1834) accepted the placement of Roth’s species in Glinus but doubted its taxonomic position in view of major morphological differences from the latter genus. Under the influence of that account, Fenzl (1836) established a separate genus for this taxon, Axonothechium Fenzl, still being uncertain about its precise taxonomic identity. Recent treatments (Gonçalves 1978; Sukhorukov and Kushunina 2015) placed Glinus trianthemoides into the synonymy of Corbichonia decumbens; this placement agrees with the material used by Fenzl and designated as neotype here.