Diplachne fusca subsp. fusca (L.) P. Beauv. ex Roem. & Schult.

Snow, Neil, Peterson, Paul M., Romaschenko, Konstantin & Simon, Bryan K., 2018, Monograph of Diplachne (Poaceae, Chloridoideae, Cynodonteae), PhytoKeys 93, pp. 1-102 : 13-18

Diplachne fusca subsp. fusca (L.) P. Beauv. ex Roem. & Schult. Figure 2

Poa malabarica L., Sp. Pl. 69. 1753, nom. utiq. rej. ( Snow and Davidse 1998). Panicum malabaricum (L.) Merr., Philipp. J. Sci. 4: 248. 1909. Ottochloa malabarica (L.) Dandy, J. Bot. 69 (2): 55. 1931. Diplachne malabarica (L.) Merrill, Bull. Torrey Bot. Club 60: 635. Leptochloa malabarica (L.) Veldkamp, Blumea 19: 64. 1971. 1933. Type. Rheede, Hort. Malab. 83, t. 45. 1703! (lectotype: designated by Merrill, Bull. Torrey Bot. Club 60: 635. 1933).

Festuca reptatrix L. Sp. Pl. (ed. 2). 1: 106. 1762. Diplachne reptatrix (L.) Druce, Bot. Arr. Brit. Pl. (ed. 2), 129. 1928. Type. Linn. Herbarium (lectotype: LINN 92.20! [left-hand specimen], designated by Snow 2000).

Bromus polystachios Forssk. Fl. Aegypt.-Arab. 23. 1775. Type. EGYPT. Alexandria, P Forsskål 1016 (holotype: C [C10001861]!).

Poa contracta Retz. Observ. Bot. 3: 11. 1783. Leptochloa contracta (Retz.) Blatt. and McCann, Sci. Monogr. Imp. Counc. Agric. Res. 5: 243. 1935. Type. INDIA. JG König s.n. (fragment, K!).

Festuca indica Retz., Observ. Bot. (Retzius) iv. 21. 1786. Diplachne indica (Retz.) Spreng., Syst. Veg. 1: 351. 1825. Tridens indicus (Retz.) Nees in Wight, Cat. Ind. Pl., 106. 1794, nom. nud. Type. INDIA. JG König s.n. (holotype: LD [1287467!]; fragment, K!; isotype: BM!). Contrary to Snow (1997a: 225), lectotypification is not needed now that the location of the holotype is known.

Triodia parviflora R. Br., Prodr. 182. 1810. Festuca brownii F. Muell., Fragm. 8: 129. 1873. Diplachne parviflora (R. Br.) Benth., Fl. Austral. 7: 620. 1878. Leptochloa parviflora (R. Br.) Verloove & Lambinon, Syst. Geogr. Pl. 76: 219. 2006. Type. AUSTRALIA. Northern Territory, Arnhem Bay, "Littora Nova Hollandiae intra tropicum", R. Brown 6254 (lectotype, here designated: BM! [BM000514993]; isolectotype: K [K000899898]!; isolectotype fragment: BRI!).

Triodia ambigua R. Br., Prodr. 183. 1810. Type. AUSTRALIA. Queensland, Keppel Bay, R Brown 6253 (lectotype, here designated, BM! [BM000514991]; isolectotype: BM!; BM fragment, BRI!).

Tridens capensis Nees, Linneana 7: 324. 1832. Uralepis capensis (Nees) Kunth, Enum. Pl. 1: 319. 1833. Diplachne capensis (Nees) Nees, Fl. Afr. Austr. 256. 1841. Triodia capensis (Nees) Th. Dur. and Schinz, Consp. Fl. Afr. 5: 877. 1895. Type. SOUTH AFRICA. "Bei Doornhoogte in der Capschen Fläche, Dec. 1824, Ecklon". No specimen is cited in the protologue; lectotypification is probably needed.

Diplachne capensis (Nees) Nees var. concinna Nees, Fl. Afr. Austr. 257. 1841. Type. SOUTH AFRICA. Ad Weltevrede in ripa Gamka..., Drège s.n. (lectotype, here designated: B ([! by Snow {1997a, but unpublished}; not found electronically online]; isolectyotypes: BM! P [P00083372], [P00083373]!; fragment: PRE!). Note: Drège or a later worker assigned this species or sheet the number 2553, which at PRE is also assigned to Diplachne livida (below).

Diplachne capensis (Nees) Nees var. minor Nees, Fl. Afr. Austr. 257. 1841. Type. SOUTH AFRICA, Drège 3900 (fragment of holotype: PRE!).

Diplachne capensis (Nees) Nees var. obscura Nees, Fl. Afr. Austr. 256. 1841. Type. SOUTH AFRICA. Circa Graaf Reynet solo Karro, Ecklon s.n.; In Herb. Reg. Berol, Mundt s.n.; Ad sinum Plettenbergbai, George s.n., (fragments syntypes [Ecklon s.n., Mundt s.n.]: PRE!)

Diplachne capensis (Nees) Nees var. prolifera Nees, Fl. Afr. Austr. 257. 1841. Type. South Africa, Ad flumen Gauritzrivier solo Karro, George s.n. (Fragment of type: PRE!)

Diplachne capensis (Nees) Nees var. prolifera subvar. minor Nees, Fl. Afr. Austr. 257. 1841. Type. South Africa. Inter Los-Tafelberg et Zwartkey, JF Drège 3900 (lectotype, here designated: P [P00083374]!)

Diplachne capensis (Nees) Nees var. pauciflora Nees, Fl. Afr. Austr. 257. 1841. Type. South Africa. In districtus Caledon montibus. Since Poaceae types of Nees were destroyed at B, selection of a neotype may be necessary.

Diplachne livida Nees, Fl. Afr. Austr. 254. 1841. Uralepis livida (Nees) Steud., Syn. Pl. Glum. 1: 248. 1855. Triodia livida (Nees) Th. Dur. and Schinz, Consp. Fl. Afr. 5: 877. 1895. Type. SOUTH AFRICA. Namaqualand, In valle praerupta montium Kamiesbergen prope Kuile alt. 3000'... JF Drège (2553?; see comment above under Diplachne capensis var. concinna ) (holotype: B!; isotypes: BM! P [P00439436], [P00439437], [P00083371]!).

Diplachne pallida Hack., Bull. Herb. Boiss. 3: 387. 1895. Type. SOUTH AFRICA. Transvaal, Boshveld, Klippan, A Rehmann 5371 (holotype: W-17607!; isotype: K [K000366665]!, fragments: PRE!, US! [US-865866]).

Leptochloa neuroglossa Eichinger, Repert. Spec. Nov. Regni Veg. Beih. 40 (1, Anhang): 75. 1930, and 264, 1931. Type. TANZANIA. Pare District, Mkomazi, Lake Manga, A Peter 41078 (holotype: B†). A neotype specimen at B (!) ("S. Pare, km 164 Sumpfgebiet Mkomasi -Mkumbara, A Peter 10497") was suggested earlier ( Snow 1997a: 227) but the specimen has not been seen recently.

Diplachne cuspidata Launert, Prod. Fl. Südwestafrika 34 (160): 221. 1970. Type. NAMIBIA. Etoscha-Pfanne, Okaukuejo, 5 Meilen westlich von Okondeka im Omuramba, 27 Feb 1963, W Giess, OH Volk, & B Bleissner 6105 (holotype: M [M0103386]!; isotype: BM [BM000514989]!).

Diplachne amboensis Roiv., Ann. Bot. Fenn. 11: 34. 1974. Type. NAMIBIA. Ovamboland, Ondonga, M. Rautanen s.n. (holotype: H! [H1057541]; isotype: H [H1003258]!; Z [Z-000017953]!).

Diplachne amboensis Roiv. var. plurinodis Roiv., Ann. Bot. Fenn. 11: 36. 1974. Type. NAMIBIA. Ovamboland, Ongandjera, Okahao, pool in valley, Soini s.n. (holotype: H! ([H1057542]).

Diplachne wahlbergii Roiv., Ann. Bot. Fenn. 11: 36. 1974. Type. SOUTH AFRICA, Caput Bonae Spei, J Wahlberg s.n. (holotype: S!; isotype: K!, S!).

Type.

PALESTINE. Ab. loco, F. Hasselquist s.n. (lectotype: LINN 92.21!), typ. cons. prop .

Description.

Plants annual or (mostly) perennial. Culms (15-)40-170 cm tall, 2-5 mm wide at base, round or somewhat flattened, ascending to erect or sometimes decumbent and rooting at nodes (and sometimes strongly stoloniferous in southern Africa), often branching; nodes glabrous; internodes to 26 cm long, soft or sometimes lignified, hollow. Leaf sheaths longer or shorter than the internodes, round or somewhat flattened, glabrous on sides and margins; ligules 4-8(-15) mm long; blades mostly 5-48 cm long, 2-6 mm wide, glabrous to moderately scabrous above and below. Panicles 15-105 cm long, 2-20 cm wide, exerted or occasionally partially inserted at maturity with 3-28 branches; the branches (1.5-) 4-20 cm long, mostly alternate along the rachis, ascending, erect, or infrequently reflexed, stiff to somewhat flexuous, minutely scabrous, axils glabrous. Spikelets 6 -14 mm long, distant to tightly imbricate; florets (4-)6-12(-14); callus glabrous or hairy; lower glumes 1.9-3.0(-4.9) mm long, ovate, scabrous on midnerve, obtuse to acute, rarely bifid; upper glumes 3.0-4.7(-5.5) mm long, ovate, scabrous on midnerve, obtuse to acute, rarely bifid; lemmas 3.0-4.7(-6.0) mm long, 3-nerved, narrowly ovate to ovate, the lateral nerves pronounced and often extending to the tips, sericeous (at least below) on lateral nerves and often on midnerve, glabrous between nerves, apex obtuse to acute or acuminate, sometimes bifid, unawned, mucronate, or awned; paleas subequal to slightly exceeding lemma, elliptic, more or less sericeous along nerves; apex obtuse to acute. Stamens 1-3; anthers 0.5-2.7 mm long (typically longer in perennials), maroon or yellow. Caryopses 1.6-2.3 mm long, 0.9-1.2 mm wide, obovate in hilar profile, transversely elliptic to depressed obovate in transverse section, hilar groove lacking, smooth, brown; pericarp weakly adnate to endosperm.

Leaf anatomy.

Midrib present (or rarely absent) in mature leaves; central lacuna present. Primary bundles: protruding adaxially and abaxially and to a greater degree than secondary bundles; outer bundle sheaths continuous adaxially, interrupted abaxially; extension cells present adaxially; adaxial cells enlarged, abaxially cells not enlarged; sclerenchymatous girders present adaxially and abaxially, or abaxially only as strands. Colourless cells present between primary and secondary bundles; chlorenchyma continuous or discontinuous between adjacent bundles. Secondary bundles: protruding adaxially or not, flush abaxially; outer bundle sheath continuous or interrupted abaxially; sclrenchymatous girders present abaxially. (Vouchers [all at MO] and country of orgin: Snow & Chatakutah 6767, 6829, 6908 (Botswana); Snow & Burgoyne 7108 (South Africa); Snow & Burgoyne 7176, 7196 (Namibia); Snow et al. 7208, 7214, 7215, 7217 (Australia); Ellis specimens from South Africa [all at PRE]: Ellis 1908, 3410, 3653, 4779.)

Stem anatomy.

Stems can root at the nodes and functionally act as stolons (e.g. Hitchock 24410 [US], Drège 3900 [P]; Valls 1978). When branching occurs, it is mostly towards the base and at lower nodes. Elliptic air canals subjacent to the epidermis of culms typically are present in populations emergent in water.

Culms hollow; pith parenchyma isodiametric; inner sclerenchymatous ring present; peripheral sclerenchymatous ring present; inner sclerenchymatous ring canal tissue present (see Culm Anatomy under D. gigantea ); Kranz sheath cells absent; Kranz sheath cell canal tissue absent. Vouchers (at MO): Snow & Chatakutah 6808 (Botswana); Snow & Chatakutah 6829 (Botswana); Snow & Burgoyne 7196 (Namibia); Snow et al. 7217 (Australia); Snow et al. 7208 (Australia); Snow et al. 7239 (Australia); Snow et al. 7249 (Australia).

Chromosome numbers.

n =10 ( Gould 1958, 1966, 1968; Reeder 1971; Valls 1978; Spies et al. 1991); 2 n =19 (Spies and Vogues 1988); 2 n =20 ( Larsen 1963; Bir and Sahni 1986).

Phenology.

Flowering throughout the year but seasonally locally.

Distribtution.

Native: Much of paleotemperate and paleotropical areas; mostly open mesic areas, often in saline conditions. Elevation sea level to ca. 2000 m. (TDWG: ANG, BOT, BZE-PB, BUR, CHA, ETH, JAW, IND, IRQ, ITA-IT, KEN, MDG, MLW, MLY-SI, MOZ, MYA, NAM, NGA, NGR, NSW-NS, NTA, OFS, PAK, PHI, QLD-QU, SEN, SOA, SRL, TAI, TAN, THA, UGA, VIC, WAU-WA, YEM-NY, ZAI.) Non-native: Persisting in Argentina, but non-persisting elsewhere. (TDWG: AGE-BA, AGE-ER, AGE-MI, AGS-RN, AGW-LR, AGW-SJ, BOL, CAL, CZE-CZ, GBR, GER, NWG-PN, SCA, SWI.). Diplachne fusca subsp. fusca was grown in experimental gardens in the early part of the 20th century in Chico and Berkeley, California, as introductions from New South Wales (McKee, SPI 17213 [NA]; Wight 1359 [NA] (Beetle s.n., 1941 [NA]). It has been collected outside the plots in California, but evidently has not become widely established in that state.

Conservation status.

Widespread; not of concern.

Etymology.

The Latin fuscus refers to dark, dusky, or swarthy and possibly alludes to the dark green or plumbeous spikelets of some specimens.

Vernacular names.

Malabar sprangletop; Beetlegrass sprangletop. Japan: Hama-gaya ( Koyama 1987). Tanzania: Msinti (Fipa and Kifipa); Milepa (Bullock 3605 and 2474, US); Hîsh (Simpson 5183, B). Iraq: Sabut (Springfield 12545, US). Egypt: Sayfoum, (Drar 9/123, NY). China: Shuang fu cao. Myanmar: Myet-cho.

Comments.

Diplachne fusca subsp. fusca is the most widely distributed taxon in the complex and occurs throughout the range of the species in the Paleotropics, with some introductions in the Neotropics and North America (California, South Carolina). It is sometimes considered a weed of rice paddies ( McIntyre and Barrett 1985, McIntyre et al. 1989).

Distinguishing D. f. subsp. fusca from subsp. Diplachne fusca fascicularis where the two are sympatric can be difficult, particularly in Argentina ( Snow 2012), where the populations of Diplachne fusca subsp. fusca may be non-native. The authors’ interpretation of taxa in the complex differs from Nicora (1995), who synonymised Diplachne uninervia (J. Presl) Parodi var. procumbens Parodi under D. fascicularis . In contrast, the authors synonymise the former under D. fucsa subsp. fusca . The subsp. Diplachne fucsa fusca is differentiated from subsp. Diplachne fucsa fascicularis primarily by the latter having panicle branches inserted at the base and having anthers shorter than 0.5 mm, whereas the former more typically has fully exserted panicles and anthers exceeding 0.5 mm (sometimes significantly so).

The floret morphology of some specimens from western Africa (Audru 2704; Adam 19242; Garba 7471; Trochain 878 [all at P]), southern Africa (Snow & Chatakuta 6908 [KSP]; Strohbach & Sheuyange BD3162 [US; Fig. 2 ) and New Guinea [Backer 16301 {duplicates at P, U, and W}]; Backer 24130 [U]; and Brass 6065 [BRI]) closely resembles that of subsp. D. f. subsp. uninervia . The specimens from New Guinea may be from New World propagules that arrived during World War II. The spikelets of the type specimen of Diplachne pallida Hack. also closely resemble those of D. f. subsp. uninervia .

The length of the awns varies throughout the range of D. fusca subsp. fusca and is too inconsistent to be of taxonomic value.

The sexuality of the florets varies widely. A population from Namibia (Snow & Burgoyne 7196 [with numerous duplicates distributed]) had male, female and hermaphroditic florets within a single spikelet. The number of stamens can likewise vary within a spikelet from one to three (e.g. Snow and Burgoyne 7196), which has been noted also by Nicora (1995). The anthers also vary considerably in length between specimens, but typically are consistent within a specimen and within populations.

Thrips (Order Thysanoptera ) of an indetermined genus were found in virtually every floret of fresh material of a specimen of D. fusca subsp. fusca from Queensland, Australia (Snow et al. 7249-F). In a study of germination, populations in Australia were affected more by nighttime than daytime temperatures ( Myers and Morgan 1989).

Some populations of this subspecies have high to very high levels of tolerance for saline or sodic soils ( Beadle 1948a - b; McVaugh 1983; Hackett and Wickens 1984; Myers and Morgan 1989; Kernick 1990; Chapman 1996; Ahmad 2009). This tendency is most developed in Australia, Asia and Africa and is least well documented for subsp. Diplachne fusca uninervia . For example, the label of Trapnell 1112 in Zambia states the species grows amidst ca. 5 cm of solid salts. Another specimen from Zambia (Greenway 6228) states that the species becomes a dominant pioneer in areas trampled out in brackish pans in grassland dominated by the grass genus Hyparrhenia E. Fourn. Others also have noted the high levels of salinity tolerance ( Reinhold et al. 1986; Hurek et al. 1987; Reinhold-Hurek et al. 1993). Joshi et al. (1983) reported salt excretion on the leaves via small apical pores in papillate structures that they designated as salt glands. Chapman (1996) also reported that some populations of D. f. subsp. fusca excrete salt from glands on the leaves and that this taxon has been advocated for its use for lowering salt levels in soils. The ability of many populations of this subspecies to tolerate and even thrive in saline soils was one of many factors contributing to the decision to recognise them at the subspecific, rather than specific rank ( Snow 1997a). In southern Africa, D. fusca subsp. fusca can be common to abundant in pans (low-lying areas that accumulate seasonal moisture but which dessicate in the dry season), where it sometimes is decumbent to geniculate and stoloniferous ( Breen et al. 1993; e.g. see Hitchcock 24410 [US]; Snow 6829 [MO]).

Herbarium labels from African collections sometimes indicate grazing, presumably by the native fauna, and others indicate good grazing for livestock. The subspecies may occur in pure or nearly pure stands in Africa. In Australia, Lazarides (1970) reported the taxon as having high palatability and being preferentially grazed, but that it typically occurs in relatively small populations. Denny (1993a, 1993b: 27) and Ahmad (2009) indicated that among grasses generally, Diplachne fusca appears to maximally combine adaptations to wetness and salinity, combined with palatability, of any species of grass. Kernick (1990: 171) linked the salinity tolerance and palatability of D. fusca and suggested that additional screening for salinity tolerant races would be worthwhile. The salinity tolerance of D. fusca subsp. fusca , high viability of its seeds ( Chapman 1996; Snow, pers. obs. in greenhouse), and high palatability of the subspecies among ruminants ( Chapman 1996; Snow pers. obs.) collectively make D. fusca an excellent candidate for reclamation of saline soils ( Haq and Khan 1971; Sandhu et al. 1981; Chapman 1996; Ahmad 2009).

A remarkable biological attribute of Diplachne fusca subsp. fusca is the presence in some Pakistani populations (where it is called Kallar grass) of a recently described nitrogen-fixing genus of bacteria, Azoarcus ( Reinhold et al. 1986, 1993; Hurek et al. 1987, 1993, 1994; Reinhold-Hurek and Hurek et al. 1997). Subsequent studies documented Azoarcus to be a nitrogen-fixing endosymbiont of many grasses, including rice ( Hurek et al. 1997; Krause et al. 2006). Given increasing problems of salinisation in some areas and the biologically and economically important properties of the Pakistani populations (nitrogen-fixing, salinity tolerance, perennial growth form and high palatability), these populations should be investigated further for their reclamation potential of saline soils, given that they also could provide useful fodder. More recent research has examined the rhizosphere for bacterial species of D. f. growing in industrial sites ( Abou-Shanab et al. 2005).

Diplachne fusca has been studied recently augmented with three experimentally introduced endophytic bacteria, as a species for use in constructed wetlands for bioremediation of wastewater from tanneries ( Ashraf et al. 2017).

Specimens examined.

Afghanistan. Maymana: 25 km SW of Maymana, Alizzi & Omar 34707 (K). Angola. Huila. Morros de Cualeque, Exell and Mendonça 2704 (BM); Distrito de Huila, Pereira de Iça, Cunhama, Gossweiler 11066 (K, US). Argentina. Buenos Aires: Pdo. Magdalena, Punta Indio, Boelcke et al. 12540 (MO); A San Vincente, Zardini 246 (LP); Vicente Lopez, Parodi 7694-1/2 (BAA); Vicente Lopez, Parodi 184 (BAA); Avellaneda, prox. al cemetario, Parodi 4774 (BAA); San Isidro, Parodi 4903 (BAA); San Vicente, Parodi 8118 (BAA, US); Palermo, Parodi 175 (BAA). Entre Rios: Dpto. Uruguay, Laguna de los Negros, Nicora 3012 (SI); Dpto. Uruguay, cerca del Parque Unzué, Burkart 25675 (BAA). Misiones: Nacanguazú, Santo Pipó, Grondona and Puccinini 3188 (UC). La Rioja: Cunta de Llanos, Nicora 18527 (B). Rio Negro: Fuerte Gral Roca, Boelcke & Serrano 3122 (BAA). San Juan: Jachal, Cabrera 18025 (LP). Australia. New South Wales: 24 km S of Moree, Beadle s.n. (NE 24646), Combanning Silo, ca. 13 km E of Temora on Stockinbingal Rd., ca. 26 km W of Stockinbingal, Snow et al. 7208 (KSP, MO, NSW); Ca. 20 km E of Griffith on Ardelthan Rd., Snow et al 7213 (MO, NSW); 9.7 km S of Darlington Point on Coleambally Rd., S of Griffith, Snow et al. 7214 (KSP, MO, MU, NSW); Marrimajeela Ck., ca. 15 km N of Boolingal on road to Ivanhoe, Snow et al. 7215 (MO, NSW); 15.1 km out of Brewarrina on Collerina Rd., Snow et al. 7216 (MO, NSW); 17.9 km out of Brewarrina on Collerina Rd., ca. 4.3 km E of Bokhara Riv., Snow et al. 7217 (MO, NSW); Ca. 60 km E of Brewrrina on road to Walgett, ca. 9 km W of turnoff to Cumborah, Snow et al. 7220 (MO, NSW); Southern end of town of Walgett, along muddy margins of artificial pond created by irrigation regime in the civic park, Snow et al. 7222 (MO, NSW); 46.6 km E of Connamble on road to Baradine, Snow et al. 7226 (MO, NSW); 19.9 km N of Boggabri on road to Narrabri, Snow et al. 7232 (MO, NSW); 6.5 km N of Edgeroi, between Narrabai and Moree, Snow et al. 7234 (MO, NSW); 11.6 km N of Moree on road to Boggabilla (Hwy 39), Snow et al. 7237 (MO, NSW); Ca. 58 km N of Moree on road to Boggabilla (Hwy 39), Snow et al. 7244 (MO, NSW); Grenfell, Corbett s.n. (BISH); Duck Ck., Wollongbar, McBarron 6516 (NSW) and McBarron 6516 (NSW); Nevertire, Helms 31318 (NA, NY, SMU, US); Walgett, Vickery 18009 (US); Macquarie Marshes area, 16 km SE of Carinda, Paijmans 1737 (CANB); Bogan Gate - 4 km E of Parker Rd., Lloyd 156 (CANB); Moree - 3 km E on Gwyndir Hwy, Lloyd 726 (CANB); 28 mi. S of Nyngan, along road between Nyngan to Narromine, Martensz 268 (CANB); Macquarie Marshes, Warren, Goodrick 261 (CANB); Agric. Res. Station, Yanco, Boerema 7 (CANB); Riverina Dist., Booroorban, between Deniliquin and Hay, Brown 390 (BRI, CANB); Near Barham, Ganba 3516 (CANB); Griffith, 1.7 km from W end of Hamilton Rd., Lodder and Corbett 11 (CANB); Near Homebush, Sydney, Hubbard 8185 (BRI); Fort Bourke, Pear Bore No. 1, Simon 2957 (BRI); 14 km SE of Bourke on Bourke-Nyngan Rd., Simon 2974 (BRI); Near Goonery Bore, 65 mi. E of Wanaaring, Moore 5781 (BRI); Between Fort Grey and Cameron’s Corner, Corrick 6536 (BRI, MEL); Macquarie Marhes 35 km W of Quambone, just E of Macquarie Riv., Paijmans 1704 (CANB); West of Premer, Hosking and Sullivan 472 (MEL, NE); Gulpa, Brown 389 (MEL); "Mt. Mulgah" - ca. 60 km NW of South, Moore 8231 (CANB); "Mt. Mulgah" - ca. 50 mi. NW of South, Moore 5091 (CANB); Ca. 51 mi. W of Bourke on Wanaaring Rd., Moore 5112 (CANB); 12 km W of Booligal, Paijmans 1649 (CANB). Northern Territory: MacDonnell Range, Palm Valley Canyon, Finke Riv. Canyon, SW of Alice Springs, Vasek 680915-25 (CANB, RSA); 40 km W of Suplejack Homestead, Latz 8121 (BRI); Gardens Station, Latz 1939 (BRI, CANB); Warangaiyu Lagoon, Elcho Island, Latz 6270 (CANB); Coomarie Spring, Latz 3956 (CANB); 7 km SW Borroloola, Latz 1523 (BRI, CANB); Homestead Ck. Bing Bong Station, Latz 1493 (CANB); Gulf of Carpenteria, Maria Island, Dunlop 2800 (BRI, CANB); Swamp below Howard Springs, Darwin area, Craven 4474 (CANB); Coastal Plains Research Station, 30 mi. SE of Darwin, Lazarides 6810 (CANB); South Alligator Floodplain, 16 km S Arnhem Hwy, Wightman 1456 (BRI, CANB); Glen of Palms, James Range, Gardner 11728 (MEL); Glen of Palms, ca. 112 km SW of Alice Springs, Beauglehole 23846 (BRI, MEL); 11 km from Borroloola, Daly Waters Rd., Jacobs 1670 (CANB, NSW); Fogg Ck. Dam near Beatrice, Jacobs 1767 (CANB); Palm Valley, 10 mi. S of Hermannsburg Mission, Perry 3501 (BRI, CANB); Mangrove area on banks of South Alligator Riv. Paijmans LAC4486 (CANB); Palm Valley, 12 mi. SW of Hermannsburg Mission, Lazardies 5290 (BRI, US); Finke Riv. 2 mi. S of Hermannsburg, Latz 3125 (BRI); Darwin, Leanyer Swamp, Latz 3616 (BRI); Fish Ck., Woolner Station, Rankin 2491 (BRI, CANB); Wirra Lagoon near Nhulumbry, Scarlett Y-257-74 (BRI); Palm Valley, Beauglehole 10359 (BRI); Palm Valley, Beauglehole 27518 (BRI); Mary Riv. Conservation Reserve, Liddle 1102 (BRI). Queensland: Burke Dist., 12 mi. E of Hughenden Township, Lazardies 3605 (BRI, MO, US); Between Mt. Emu Plains and Mt. Sturgeon Stations, Hubbard & Winders 7560 (BRI); Normanton, Blake 9002 (BRI); Agnes Lake, 79 km S of Lyndhurst Station on Hann Hwy, Simon and Clarkson 2752 (BRI); Mount Isa, Winders 7427 (BRI); Nonda, Hubbard & Winders 7228 (B, BRI, CANB, K, MO, NSW); Mt. Isa, No. 7 Tailings Dam, Schmid 712 (BRI); 3 km NE of Burketwon on Truganini Crossing Rd., Jacobs 1318 (CANB, NSW); Dry bed of Lake Louisa ca. 90 mi. N of Hugenden, Blake 8605 (CANB); 19 mi. S of Mt. Isa Township, Lazarides 4381 (US); 18 mi. SW of Claraville Station, Lazarides 3934 (CANB); 8 mi. E of Iffley Station, Lazarides 3939 (CANB); 8 mi. N of Cloncurry Township, Lazarides 4315 (CANB); 3 mi. SW of Malbon Township, Lazarides 4400 (CANB, MO). Cook Dist., 1.6 km S of mouth of North Kennedy Riv. on Lakefield N.P., Neldner and Clarkson 3782 (BRI, NSW); Arriga area, ca. 13 km SW of Marreba, T. Mailsel’s Farm, Pregno s.n. (BRI); 2.5 km SW of the aboriginal settlement at Mapoon, Clarkson 4962 (BRI, K); Weipa, Morton 1132 (MEL); 10 mi. NNE of Lyndhurst Station, Lazarides 3737 (CANB); Station Ck. saltpan, 1 km E of Inkerman homestead, Neldner & Clarkson 2873 (BRI, NSW); On southern bank of Nassau Riv., 0.5 km upstream from junction with Rocky Ck., Neldner & Clarkson 3007 (BRI); 9 km NW of Kenchering by road, Dalliston CC437 (BRI); Chohan’s Farm, Ariga W of Mareeba, Hawton s.n. (BRI); Lakefield N.P., Knifehole, 2 km from Saltwater Ck. crossing on the Musgrave to Lakefield Rd., Clarkson & Simon 7061 (BRI); Plains on eastern bank of Nomenade Ck., Morton AM1132 (BRI); 2 km N of the Morehead Riv. Crossing on the Lakfield to Musgrave Rd., Clarkson & Simon 7047 (BRI); 3 mi. ESE of Spring Ck. Station, Lazarides 3807 (BRI). Darling Downs Dist., ca. 10 mi. SW of The Gums on roadside, Johnson 729 (BRI); Kindon Station, 54 mi. NNW Goondiwindi, Smith 549 (GH); 28 km E of Goondiwindi on road to Yelarbon, Inglewood and Warick, Snow et al. 7249 (BRI, KSP, MO, NSW); 3.2 km N of junction with Cunningham Hwy (Hwy 42) on the Leichhardt Hwy, N of Goondiwindi, Snow & Simon 7312 (BRI, MO); 14.5 km N of Cunningham Hwy along the Leichhardt Hwy N of Goondiwindi, 3.1 km S of turnoff to Toowoomba (78 km S of Moonie), Snow & Simon 7326 (BRI, MO); 17.8 km W of Moonie on Moonie Hwy, Snow & Simon 7360 (BRI, MO); 22 km W of Moonie along Moonie Hwy, Snow & Simon 7363 (BRI, MO); Palardo, W of Miles, Blake 7619 (BRI, CANB, NSW); ca. 10 mi. SW of The Gums, Johnson 729 (CANB, NY); Yelarbon, Blake 10445 (BRI); Yelarbon 41 km E of Goondiwindi, McDonald 406 (BRI, CANB, K); Chinchilla, Hubbard and Winders 6405 (B, BRI, CANB, MO); ca. 7 mi. SE of Meandarra, Johnson 1206 (B, BRI); Condamine State School, Turnbull 5 (BRI); Milmerran, Hubbard 5867 (BRI, CANB); Dalby, Beiers 43 (BRI); Chinchilla, Beasley 124 (BRI); Dalby, Beiers 63 (BRI); 10 Meilen von Meandarra, Walter and Walter 2610 (B); near Gurulmundi, Hubbard 5129 (B, CANB, MO); Yelarbon, Everist 889 (BRI); Pelican Lakes 30 km NE of Chinchilla, Simon et al. 3491 (BRI). Gregory North Dist., Boundary Gully Bore, 25 km N of Headingly homestead, Neldner & Stanley 1804 (BRI). Gregory South Dist., 0.7 km SW of Longford turn-off, Thompson Devel. Rd., Prendergast HDP313 (BRI); ca. 92 km W of Betoota, Nicolson & Novelly 277 (BRI). Leichhardt Dist., Broadsound Shire, “Iffley”, 3 km N of Mt. Coxendean, Anderson and Russell 896 (BRI); Broadsound Shire, Anderson 752 (BRI); Emerald, Finlay & Farquhar 2 (BRI); Broadsound Shire: "Barwon Park" ca. 70 km N of Blackwater, Anderson 2054 (BRI) and (ibid.) Anderson 2055 (BRI); Gunnewin via Roma, Kieseker 9 (BRI). Maranoa Dist., Noondoo near Dirranbandi, Blake 10690 (BRI); 13 km E of Mervyndale, Walrus III, Site R030, Purdie 462D (BRI); 40 mi. W of St. George, Roe s.n. (CANB); “Warrie”, Nindigully, Allen 111 (CANB); Noondoo Station E of Dirrabandi, Everist 817 (BRI); Mitchell, Hubbard and Winders 6309 (BRI); Boatman Station, Everist 2857 (BRI, K, NY); 15 km S of Roma, Bungil Shire, Silcock S721 (BRI); Roma, Blake 10894 (BRI); Bollon, along damp creek bands, White 11545 (GH, US). Mitchell Dist., Yaraka, Bownman s.n. (BRI); 2 mi. W of Blackall, Everist 2089 (BRI); 34 mi. NW of Longreach, Davidson 82 (BRI); 11 km SE of Corinda homestead, Thompson and Sharpe MUT1 (BRI); Jericho, Blake 10245 (BRI). Moreton Dist., Wyampa, near Bald Hills, Blake 213 (BRI); Near Redcliffe, Hubbard 5551 (B, BRI, CANB, NSW); Currumbin, White 8728 (BRI, NY); Nundah, Brisbane, Blake 159 (BRI); 2 km N of Coolum beach, ca. 130 km N of Brisbane, Sharpe 2127 (BRI); Serpentine Ck. and environs, ca. 11 km NE of Brisbane, Durrington 435 (BRI); 2 km N of Coolum Beach, ca. 130 km N of Brisbane, Sharpe 1894 (BRI); Hope Island, 2 km from Boyankil on road to Pacific Hwy, Simon et al. 2932 (BRI). North Kennedy Dist., Toonpan - Antil Plains, Everist 9194 (BRI); Mt. St. John, Hubbard & Winders 6943 (BRI, CANB); Barratta Ck., lower Burdekin Valley, Paijmans 3508 (CANB); Near Saltern Lagoon, 5 km W of Valley of Lagoona HS, Lazarides 8167 (BRI, CANB); Cromarty, between Townsville and Ayr, Blake 8306 (BRI). Port Curtis Dist., Gladstone, Blake 12784 (BRI); Curtis Island, N of Southend, Blake 22578 (BRI, CANB, MO); Curtis Island, S end, Blake 22522 (BRI); 25 km ESE of Rockhampton, Paijmans 2016 (CANB); E side of Herbert Ck., 5 km NW of “Banksia” homestead, Paijmans 2017 (CANB); Gladstone, Blake 12784 (CANB); “Torilla” between Broad Sound and Shoalwater Bay, Blake & Webb 15615 (BRI, NSW). South Kennedy Dist., 17 mi. NE of “Mirtna” Station, Adams 1133 (CANB); `Cassiopeia’, the property of J. Mendazona, Belyando Shire, Jacobsen E 580 (BRI); 6.5 km NW of Belyando Crossing, Thompson and Sharpe BUC901 (BRI); Gormans Lagoon near Mirtna HS (site GL1), Thompson & Simon BUC755 (BRI); with 2.3 km S of Doongmabulla homestead, 197 km N of Jericho, Thompson & Henderson GAL142 (BRI); 10.5 km NE of Hyde Park homestead on road to Bulliwallah homestead, Thompson & Simon BUC813 (CANB, K); 3 km S of Doongmabulla homestead on sandy flat adjacent to Cattle Ck. [...] Thompson & Simon GAL50 (BRI); Laglan Station, ca. 105 km WNW-NW of Clermont, Smith 10293 (BRI); 3 km W of Yarromere homestead, Thompson & Henderson BUC993 (BRI). Warrego Dist., ca. 25 mi. S of Wyandra, Blake 11234 (BRI, K); Wyandra, Sutton s.n. (BRI); Curragh Station, Hubbard & Winders 6263 (K, MO, NSW); Gilruth Plains Stn, Cunnamulla, Barker 835 (CANB); “Cowley”, 40 mi. E of Quilpie, on main Quilpie-Charleville Rd., Anson 2 (BRI); Gilruth Plains, Cunnamulla, McKee 10326 (BRI, NSW); 10 km SE of Charleville along Boatman Rd., Purdie & Boyland 36 (BRI); Coongoola, between Charleville and Cunnamulla, Hubbard & Winders 6174 (BRI, CANB); “Ambathala”, 100 km WNW of Charleville on Adavale Rd., Silcock 5458 (BRI); “Peneroo” via Eulo, Young 54 (BRI); Bowalli, ca. 75 mi. SSW of Quilpie, Everist 5041 (BRI); “Wittenburra” Eulo, Ebersohn E180 (BRI); Carbeen near Cunnamulla, White 11567 (BRI, GH); Dynevor Lakes, 30 mi. E of Thargomindah, Blake 11761 (BRI); Bulloo, MacGillivray 1024 (BRI). Wide Bay Dist., Splitters Ck., Smith 437 (BRI). South Australia: 30 km W of Inllian Ck on Coober Pedy road, Badman 1282 (AD, BISH); Lower Winkie Rd.... near Berri, ca. 195 km NE of Adelaide, Kuchel 3215 (AD, BISH, BRI, MO); 32 km SW of Hawker Gate, Jacobs 3536 (BRI); Just out of Wyndham on and at base of rocky slope, Symon 5251 (CANB). Victoria: Melbourne area, between Newmarket and Showgrounds railway stations, E of Ascot Vale Rd., Flemington, Clarke 1797 (BRI, MEL); King’s Billabong, beside the Murray Riv., between Mildura and Red Cliffs, Henshall s.n. (MEL); Murray Valley area, Tatura - Irrigation Research Institute, Brown 34 (MEL); Port MacKay, Dietrich 538 (MEL); Malle Study Area, Reedy Lake Wildlife Reserve, Beauglehole 55705 (MEL). Western Australia: Wittenoom Gorge, Hammersley Range, Beauglehole 11564 (BRI); 10 km SE of Wyndham, Paijmans 2505 (CANB); 8 km SE of New Wyndham Post Office, North Kimberely, George 14546 (CANB, K, NSW); Goody Goody, Fitzgerald 65 (US); 34 mi. SW of Wyndham Township, Lazarides 3080 (NSW, US); Wyampa, near Bald Hills, Blake 213 (K); Parry Lagoon, NE Kimberley, Kenneally 10719 (CANB); Ord Riv. 25 km NE of Wyndham, Paijmans 2268 (CANB); North West Coastal Hwy, 94 km from Minilya on raod to Dampier, Simon & Stretch 3788 (BRI); 34 mi. SW of Wyndham Township, Lazarides 3080 (BRI). Belgium. Graviers de la Vesdre, Fasseaux 11a (K). Bolivia. Beni: Prov. Ballivian, Espiritu en la zona del influencia del rio Yacuma, Beck 3219 (US). Santa Cruz: Chiquitos, Est. San Ignacio, 22 km N of San Jose de Chiquitosm Killeen 1712 (MO, US); Nuflo de Chavez, San Antonio de Lomerio, Killeen 825 (MO, W). Botswana. Central: Mumpswe Pan, 25 mi. NNW of mouth of Nata Riv., Drummond & Seagrey 5167 (PRE); 17.6 road km N of Dibete (at turnoff to Mookane), ca. 135 Road km N of Gabarone, Snow & Chatakuta 6767 (GAB, K, MO, PRE); At km marker 16, W of Nata on Hwy to Maun, Snow & Chatakuta 6829 (GAB, K, MO, PRE); Along main Hwy between Nata and Maun, ca. 67 km W of Nata, Snow & Chatakuta 6878 (GAB, K, KSP, MO, MU, PRE); 44.3 road km from Sephophe on road towards Zanzibar, ca. 21.6 km prior to Tsetsejwe, Snow & Chatakuta 6908 (GAB, K, KSP, MO, MU, PRE); Along main hwy, 17.6 km N of Dibete (at turnoff to Mookane), ca. 135 road km N of Gaborone, Snow & Chatakuta 6913 (GAB, K, MO, PRE); Botletle delta area, NE of Mopipi, Tyers 616 (MO). Kgalagadi: Zanye Pan, NW of Hukuntsi, Ellis 2657 (K, MO, PRE). Mahalapye: Kalamare, de Beer K5 (GAB). Mohembo: Okavango Riv., Smith 2669 (K); in the sump of Nwaku Pan, Smith 2969(K). Northern: Mumpswe Pan, 25 mi. NNW of mouth of Nata Riv., Drummond & Seagriest 5167 (GAB, K); Kangwa (Xanwe), 27 km NE of Aha Hills, Wild & Drummond 6931 (BM, GAB, K); near Tschelenyane, 11 km from Lake Ngami on track to Khwebe Hills, Drummond 8747 (K). Ngamiland: Boteti Riv. lower reaches, Smith 2563 (K, PRE, US); Nxai Pan N.P., Smith 1648 (K, MO); Moremi Wildlife Reserve, Smith 811 (GAB, K, MO, PRE); Moremi Wildlife Reserve, Mexara Pan, Smith 1598 (BRI, K). Brazil. Paraiba: São João do Carirí, Pôsto Agro-Pequário, Mattos and Mattos 9736 (US). Burundi. Plaine de la Rusizi au N de Bujubura, Gihanga, Van der Veken 11182 (B, U); Plaine de la Rusizi (Prov. Bubanza), savane-palmeraie a hauteur du km 14 de la route Bujumbura-Cibitoke, Lambinon 75/42 (M, MO); Vallée Katunguru, Reekmans 3187 (MO). Chad. Entre Moolou et Berirem, Chevalier 10102 (P); Latir Nord Bol, Dune, Gaston 749 (P); Bahr el Ghazal-Gahine, Zolotarevsky et al. 876 (P); Bagana - Ba Gara, Murat 924 (P). China. Hebei: Peitaiha [=Beidaihe], Cowdry 213 (US); Tientsin, Clemens 1615 (B, BM, MO); Shen Hsien, Beach s.n. (US); Shatin, Shiu Ying Hu 12084 (A). Hong Kong: Hong Kong, Shiu Ying Hu 11785 (A, CANB). Hopei: Ch’ating, Kung 3784 (US). Heijan: Roadside ca. 10 mi. W of Hochien, Beach s.n. (US). Jiangsu: Nanking (Lotus Lake), Reeves s.n. (NY). Democratic Republic of the Congo. Sud-Kivu: Plaine Rusisi [=Rusizi], Germain 5576 (B, M). Distr. Liberec: garden in Raspenava, introduced with waste from wool processing, 24 Aug. 1963, V. Jehlík s.n. (PRA). Egypt. [Province Unknown: Lake Manzala: Masraf Abu Aziza, Mustafa & Sabet s.n. ( KSC 77803 )] Damietta District, Ezbet El-Burg, Mashaly s.n. (K); Aus der Umgegend von Cairo, Schweinfurth 1432 (K); Al-Faiyum District, El-Azab, Ghani 3650 (K); Faraskûr, Shabetai 33 (K); Aulad Hammam, Simpson 5183 (B); Gize, Kairo, Kneucker 865 (MU); near Cairo, Meinertzhagen, s.n. (BM); Fayoum, Drar 9/123 (NY); 166 km from Cairo on the desert road to Alexandria, Amin et al. s.n. (NY); An den Rändern der Bewässerungsgräben zwischen dem Dorfe Gîze unweit Kairo, Kneucker 254 (PR, US); Between Hamul and Baltin, Hefnawy s.n. (K); Lake-side, Manzale N of Damietta, Simpson 1451 (K); Wâdi el Gedîol-Dâkhlao, E part of the oasis below Balat and Ismant (Smint), Walter s.n. (K). Ethiopia. Oromia: Lake Alemaya (Haramaia), Burger 3607 (PRE, US); South shore of Lake Alemaya, 35 km on the road from Dire Dawa to Harar, de Wilde 4789 (B, MO); NW shore of Lake Langano, Ash 1377(K, MO) and 2644 (K); Lake Awasa, ca. 20 km SW of Shashamane, de Wilde & de Wilde-Duyfjes 7016 (B, MO). Germany. Wollkammerei, Kuhbier s.n. (B). Indonesia. Java: Horsfield s.n. (BM). India. Kerala: Vembanad Lake near Alleppey, Vencoba Rao 4061 (K). West Bengal: Salt Lake, Calcutta (Kolkata), Clarke 21605 (BM). Iraq. Basrah: Basra [h], Omar & Omar 35105 (K); Ambadi Basra, Graham 535 (BM); Garma inter Basra et Qurna, Rechinger 15815 (B, K, M, S, UC, US, W). Al-Chibayish: Al-Chibayish [as Chabaish], Thamer 50106 (K). Unknown: Maqil, Rechinger 8451 (M, W); Kabajish, Springfield 12545 (US). Italy. Venice: Venice, Eggert s.n. (NY, KSC). Kenya. Coast: Malindi, Banks of Sabaki Riv., Bogdan 2605 (UC). Rift Valley: Mile 5 from the Sobo Rd.-Galana Riv. turnoff, Greenway & Kanuri 12759 (P); West Shore of Lake Baringo, Bogdan 4102 (P, US); W side of Lake Baringo, Burney & Burney B66 (NY); Masai Distr: N end of Lake Natron, SW of Shombole Mt., Milne-Redhead & Taylor 7010 (B, K). Madagascar. Mahajanga: Ambohimena (Marovoay, Ouest), Bosser 8357 (P); Marovoay, Perrier de la Bâthie 11216 (P). Toliara: Antanimora (Pce. du Fort-Dauphin), Decary 4636 (P); Vohitany, Bosser 15614 (P); Vohitany, Bekily, Bosser 15686 (P); Ambovombe, Decary s.n. (P); Beloha-Ranolava (Tsihombe), Zolotarevsky 2623-37 (P). Malawi. Polambe plain near Rjalo Island, Jackson 582 (K, US); Dwangwa Riv., Jackson 1111 (K); Fort Johnston, Jackson s.n. (P). Mozambique. Inhambane: San Sebastian, Mogg 29142 (PRE). Maputo: Maputo [as Lourenco Marques], entre a Costa do Sol e Vila Luiza, a 5 km da vila, Fidalgo de Carvalho 1446 (MO); Maputo [as L. Marques], estrada para a Costa do Sol, Balsinhas 656 (B); Inhaca Island, 23 mi. E of Lourenco Marques, Mogg 29850 (PRE); Inhaca Island, Mogg 29799 (MO). Sofala: Beira, Hitchcock 24410 (US). Myanmar. Bago: Bago [as “Pegu”], Kurz 1105 (K). Namibia. Caprivi: Asheshe area, ca. 2 km S of Asheshwe village, Ward 36 (PRE). Hardap: Gibeon: 63 km N of Tses along Marietal-Keetmanshhop Hwy, Davidse & Loxton 6342 (BRI, MO, PRE, US); Farm Helmer inghausen, Kinges 2198 (PRE); NNW of Maltahöhe on Rd. D850 to Büllspoort [Bullsport], Smook 9338 (US). Karas: Along Hwy B4, ca. 62 Road km E of Goageb, ca. 40 Road km SW of Keetmanshoop, along Fish Riv., ca. 100 m N of bridge over river, just downstream from cement irrigation channel over river, Snow & Burgoyne 7176 (K, KSP, MO, PRE, WIND); Hardap Region, 34.4 road km N on C14 from junction of C14/C13, between Helmeringhausen and Maltahöhe, Snow & Burgoyne 7196 (K, KSP, MO, PRE, WIND); 27 km W of Maltahohe on Rd. no. 36 to Sesriem, Ellis 4779 (PRE). Kohmas: Neudam Exp. Farm, Van Vuuren 1033 (K, PRE); In vlei at Ludwein near Windhoek, Liebenberg 5074 (UC). Kunene: Etosha Game Park, 11 mi. N of Okandeka waterhold, Tinley 1202 (K); Etosha N.P., 8 km nürdlich von Okaukuejo am Weg nach Okondeka, Giess & Müller 13960 (M, PRE); Estosha Game Park border, Ekuma Riv., Tinley 1140 (K). Ohangwena: Ovamboland Nature Reserve, de Winter & Giess 6916 (K) Okavango: Klein Omuramba 5 mi. E of Runth, de Winter 4076 (K). Omaheke: Farm Omupanda, 8 km E of Hochveld, Gibbs Russell & Smook 5356 (K) and 5357 (PRE). Oshana: Ondangua, Soini s.n. (H). Oshikoto: Oniipa, Soini s.n. (H). Otjozondjupa: Sonop Research Stn, Strohbach 2309 (K); 35.3 mi. SW of Taumeb on road to Otavi, de Winter 2916 (K); 27 km W of Maltahohe on Rd. no. 36 to Sesriem, Ellis 4779 (PRE); Farm Tjo Noord, 25 km E of Kalkveld, Gibbs Russell and Smook 5248 (K); Waterberg Omuramba, Volk 1420 (US). Region unknown: Ab loco, Felmer 456 (US). Niger. Zinder: Zinder, Hagerup 576 (GH, S). Region unknown: Point Sabonjari, Boudet 5040 (P). Nigeria. Bornu: North Eastern State, Baga Lake Chad Border, Wit et al. 1474 (MO); N of Mardufuri, de Leeun 1896 (P). Pakistan. Punjab: ca. 10 mi. from Lahore on way to Baidyan, Qaiser 3551 (K, TAES). Papua New Guinea. Western: Mainland, opposite Daru Island, Brass 6065 (BM, GH, NY, US); Near Bula Village, at mouth of Morehead Riv., Pullen 7029 (A, K, US); near Mabaduan Hill, 50 km WSW of Daru, Paijmans 1501 (CANB). Philippines. Luzon: Manila, Merrill 554a (B, BM, K, PR, UC, US); Hot Springs, Los Baños, Gates 6348 (KSC); Los Baños, Merrill 5104 (K, US, W); Los Baños, Williams 2027 (K, NY, US); Manila, Santos 27 (US); Barrio Tanuk, Buluan Marsh, SE bank of Buluan Lake, Vera Santos 5969 (B, US). Pampanga: Masantol, "Fishery Student" s.n. (PNH). Saudi Arabia [provinces unknown]; km 106, Makkah by-pass, Collenette 3938 (K); Al-Taif, Al-Hada region, ca. 20 km from Al-Taif, Fayed 1151 (K). Senegal. Dakar: Hann, Adam 14090 (MO); Rufisque, Rte de Mbour, Adam 2273 (MO); Dakar Bongo, Audru 2783 (P); Rufisque, Route de M’Bour, Adam 2273 (MO). Kaolak: Ravin des Voleurs, Berhaut 3784 (P); Kaolak, Trochain 4035 (P); Kaolak, Berhaut 626 (P). Louga: Menguilé, Ferlo, Mosnier 2405 (P). Saint-Louis: St. Louis, Berhaut 2813 (P); St. Louis, Khor Marshes, Hepper 3613 (P); Tiguet, Audru 2938 (P); N’Diael, Audru 2982 (P). Unknown: Mbidjem, rive W du Tanma, Raynal 6559 (P). Singapore. Perak, Ag. Officer 35646 (BM). South Africa. Cape Districts (Northern, Western, Eastern): Beaufort West Distr., Farm Layton, Shearing 267 (K); Dist. Swellendam, Nat. Bontebok Park, Liebenberg 7204 (K); Near Riversdale, Muir 2859 (K); Gordonia, Kalahri Bemgsbok N.P., Leistner 1017 (K); Ft. Beaufort, between Ulster and Mooiriver, Smook 4033 (K); Dist. East London, Bulura Mouth, Eacocks 15785 (K); Dunswart, near Johannesburg, Moss 13959 (K); Duivenhoksrivier, 10 km NW of Vermaaklikheid on road to Heidelberg, Davidse 33779A (MO); Springbok Dist., Namaqualand, top of Wildeperdehoek Pass SW of Springbok, Goldblatt 2820 (MO); Piquetberg, Liedenberg 4266 (UC); Upington, Gordonia Distr., Liebenberg 4163 (P, UC); Buffelsrivier valley between Pedroskloof and Bobbejaanhoek on road to Rooifontein, Davidse 33309 (MO); 10 km S of Humansdorp on road to Cape St. Francis, Davidse 33626 (MO); NW...on Holbakrivier in small port, Smook 3858 (K, US); Carnarvon, Acock 1737 (US); Cape of Good Hope at Douglas Heights, NW of Grahamstown, Godfrey and Storey SH-1362 (US); Hardenbeck Riv. dam cistern on Excelsior Farm, Hugo 2286 (MO, US); Farm Lemoenkop 337, ca. 2 km N of Lemoenkop farmhouse, just S of True Beacon 25 in drainage channel, Le Roux & Lloyd 94 (PRE); Nat. Kalahari Gembsbokpark, Liebenberg 7080 (PRE); Obobogorap, ca. 120 mi. NW of Upington, Leistner 1781 (PRE); Nat. Bontebok Park, Liebenberg 7204 (PRE); Kleinemonde, Ward 8897 (PRE); (Ubombo) Lower Mkuze floodplain, Ward 8785 (K); Paterson, 2 km off main road to Addo, Smook 3768 (K, PRE); 53 km SE of Graaff Reinet on small running stream below homestead, Smook 3896 (K, PRE); Karoo N.P., Doornhoek, Bengis 462 (PRE); Grootfontein, Theron 701 (PRE); Beaufort West Distr., Farm Layton, Shearing 380 (PRE); Kalahari Gembsbok Park, Mata-Mata, Van Rooyen and Bredenkamp 115 (CM); Kaffraria, Burtt-Davy 12786 (PRE). Free State: 8 km from Luckhoff on road to Jacobsdal, Smook & Gibbs Russell 2456 (K); Boskop (363) langs Petrusburg-Boshofpad, Muller 1304 (K); Small pan just S of Mayhem Pan, Edwards 4159 (K); 11 km from Petrusburg on road to Boshof, Smook & Gibbs Russell 2498 (K, PRE, US); 23 km from Petrusburg on road to Boshof at Modder Riv. Bridge, Smook & Gibbs Russell 2501 (K, PRE, US); 5 km from Bultfontein on road to Brandfort, Ellis 3653 (PRE); Bloemfontein, du Preez 1884 (PRE); ca. 10 km W of Odendaalsrus, Smook 6501 (BRI); 25 km from Luckhoff on road to Phillipolis, Smook 2879 (PRE); 5 km from Bultfontein on road to Brandfort, Smook 2728a (PRE); Distr. Harrismith, Mont Pelaan, Ferreira F212 (P); Mont Pelaan, Harrismith, Ferreira 212 (P). Gauteng: Benoni, Bradfield T371 (PRE). Kwazulu-Natal: Bridge over St Lucia Estuary, Ellis 3410 (PRE); St. Lucia Lake, Hlabisa, Zululand, Liebenberg 5903 (UC); South Coast, Pole-Evans 778 (PRE); Bridge over St Lucia Estuary, Ellis 3410 (PRE); Falce Bay Park, Ward 7724 (PRE); Bartlow Combine bay Hluhluwe Wildtuin, 24 km van Mkuzi op pad na Mtubatuba, draai na Weste vir 20 km, du Toit 660 (PRE). Limpopo: Kruger N.P., Malongafontein, Ellis 1908 (PRE); 13 km SE of Chrissiesmeer, Smook 4905 (K); On Farm Mosdene, Distr. Naboomspoint, Liebenberg 4424 (UC); Dist. Bloemhof, SA Lombard Nature Reserve, Leistner 78 (K); Kruger N.P., Malongafontein, Davidse 5875 (BRI, K); Kruger N.P., Malongafontein, Ellis 1908 (PRE); Kruger N.P., Shingwidzi rest camp, Oakes 1455 (US); Kruger N.P. 19 mi. NE of Skukuza, De Winter and Codd 560 (BM, K); Kruger N.P., near Malonga Spring, Oakes 1483 (US); Wanetzi, Van der Schyff 508 (PRE); 23 mi. SE of Punda Maria, De Winter & Codd 674 (BM, K, PRE); Mosdene farm Naboomspruit [Mookgophong], Germishuizen 42 (PRE); Naboomspruit, Schweickerdt 1791 (BM); 2 mi. N of Rust der Winter Dam, De Winter & Codd 248 (BM, CANB). Mpuma langa: 3 km from Lake Chrissie near the shore of the lake, Loxton 398 (PRE). North West: Beneath spillway of small (ca. 5 m tall) earthen dam, located below N side of road ca. 50 m, on N14 Hwy, 4.1 road km E of Kuruman, Snow & Burgoyne 7108 (K, MO, PRE); 11 mi. NNE of Lichtenbrug on road to Koster, Scheepers 1495 (K, PRE); W of Biessiesvlie, Farm Holfontein, Smook 6628 (PRE, RSA); Knopfontein 101 IP, near Coligny, Pan no. 7, Allan 117 (PRE); Knopfontein 101 IP, Pan no. 7, Allan 134 (BRI, PRE); Bloemhof, Lombard Nature Reserve, Gemsbokpan, Leistner 2108 (K, M, PRE); Bloemhoef, Lombard Nature Reserve, Jeffers 400 (PRE), Lombard Nature Reserve, Oeistner 78 (BM). Unknown: Malathlopanga, Van der Schyff 5658 (M, K, PRE); W of Biessiesvlei, Farm Holfontein, Smook 6228 (PRE, US). Sri Lanka. Jaffna: Near Ampan, Clayton 5236 (CANB, K, TAES, US); ca. 4 mi. NW of Jaffna, along the coastal road, Davidse & Sumithraarachchi 9119 (BRI, CANB, K, MO, TAES, US); Keerimalai to Point Pedro, Clayton 5201 (CANB, K, TAES, US); Near Chavakachcheri, Clayton 5255 (CANB, K, TAES, US). Mannar: Mantai, Davidse & Sumithraarachchi 9174 (BRI, CANB, K, MO, NY, TAES, US). Puttalam: Puttalam, sea front, Clayton 5664 (CANB, K, TAES, US); Puttalam, sea front, Clayton 5665 (US); Trincomalee Dist., ca. 5 mi. due S of Tamaivillu, Davidse 7581 (CANB, US). Province Unknown: ca. 4 mi. NE of Hambantota, marker 154/4, Gould 13458 (US). Switzerland. Skåne, Lackalänge, Furuland, Blom s.n. (US). Taiwan. Little Quemoy, Chuang 4459 (UC, US). Tanzania. Arusha: Sanya Plain, Arusha-Moshi, Leippert 6455 (M); Ngorogoro Crater, Greenway & Kanuri 12535 (P); Ngorogoro Crater floor, Gorigor swamp, western end, Raynal 19521 (P); Ngorogoro Crater, Heady 1517 (UC); Tituski Riv. near Lake Magadi, Greenway & Turner 10043 (US); Nyumba ya Munga Lake, at Magadini fishing village, Mhoro & Backéus 2025 (MO). Mara: Musoma Distr., Mbalageti Riv., Greenway 9027 (B, PRE). Mbeya: Ikoga, 100 mi. SW of Iringo, Heady 1871 (MO, UC). Rukwa: Lake Rukwa, Bullock 3439 (B); Near Kampunda, Bullock 3605 (US). Tanga: Mkomazi Valley, Semsei 3964 (P). Province Unknown: Lake Rukwa Extension, Michelmore 738 (US); Milepa, Lake Rukwa, Michelmore 1418 (BM, MO); SW Serengeti Plains, headwaters of the Mbalangeti Riv., Greenway et al. 13183 (K, MO); Northern Province, Scout 618 (NY). Thailand. Paknam, Sørensen et al 2038 (A, B, K, P); Ab. loc., Kerr 16107 (BM, K); Hua Hiss, Kerr 16126 (BM, K); Tachin, Kerr 8975a (K); Palanam, Kerr 20441 (K, US); Paknam, Sørensen et al. 98 (K); Paknam, Sørensen et al. 2035 (B, K); Tacheen, near Bankok, Marcan 1812 (BM); Kao Tao, Kerr 16183 (BM, K). Uganda. Butiaba, Lake Albert, Greenway & Eggeling 7047 (K). United Kingdom. Barming, Mason J/67/K1 (K); Blackmoor, N. Hants, Lousley 1348 (BM, K); Charlton, Webster 8899 (BM, K) and Webster 2053 (K); Blackmoor fruit farm, grown on at Ware, Herts, Hanson 162 (BM); Blackmoor, Wurzell 1086 (MO) and Lousley 2914 (K) and Lousley 3008 (K) and Webster 7005 (K). United States of America. California: Kern Co., Isabella Lake bed, old Kern Riv. channel ca. 1/ 4 mi. S of of the old Borel Canal Bridge, Twisselmann 6491 (MO). Santa Barbara Co., San Jose Ck. at Southern Pacific RR crossing, Goleta, Pollard s.n. (MO). Yolo Co., 2 mi. SW of Knight’s Landing, Rose 69041 (FSU, GH, MICH, MO, NY, RSA, SMU). South Carolina: Berkeley Co., Santee Wool Combing Mill, Jameston on SC Rte 45, Ahles & Haesloop 53442 (NCU); ibid., Ahles & Haesloop 47048 (NCU); ibid., Ahles & Haesloop 52773 (NCU); ibid., Ahles & Haesloop 43027 (NCU). Florence Co., Wellman Wool Combing Mill, N of Johnsonville on SC Rt. 41, Ahles 42895 (MO, USCH); ibid., Ahles 42984 (NCU), Ahles & Haesloop 30873 (NCU), and Ahles & Haesloop 30876 (NCU), and Ahles 42895 (OKL). Yemen. Rada, Westinga 3 (K); just N of Rada, Wood 3207 (BM, K). Uruguay. San José: Rio San José, Puerto Tres Bocas, Rosengurtt B-7316 (P). Zambia. Kabwe: Kakumbasa Riv., Verboom 3185 (K). Senanga: Barotseland, Mulonga and Siloana plains, Verboom 1125 (BM, K). Districts unconfirmed: Kafue flats, Magabuka, Astle 1404 (K) and Greenway 6228 (K); Monze, near Lochinvar, Rensburg 2707 (K); Lochinvar estates, Trapnell 1112 (K); Fort Jameson, Van Rensburg 2135 (B); Kafue N.P., Mitchell 24/42 (B, BM); Muckle Neuk, 12 mi. N of Choma, Robinson 587 (M); 12 mi. N of Choma, Robinson 2857 (M, P); Njeju Game Reserve, Verboom 749 (BM, K); Kakumbasa Riv., Kabwe rural, Verboom 3185 (MO); Lake Chisi, E of Mwera wa Ntipa, Michelmore 428 (K). Zimbabwe. Gwampa Forest Reserve, Goldsmith 10/56 (B, BRI, CANB, NY, P, UC, US); Lower Sabi District, Rattray 1235 (BRI, P, US); Shashi Irrigation Scheme, 12 mi. SE of Tuli, Drummond 5924 (BM); Sentinel Ranch, some 12 mi. N of Pazhi-Limpopo confluence, Drummond 5982 (BM); Beaufort West, Bleak House Farm, Gibbs Russell et al. 370 (MO); Ft. Beaufort, 3226 DD Alice, “Woodstock” H. E. Matthews’ farm, Giffen 1624 (MO).

Figures

Figure 2. A-G Diplachne fusca (L.) Kunth subsp. fusca A habit B inflorescence C ligule D spikelet E lemma F floret and rachilla G palea, lodicules, ovary, and stamens. Drawings from Strohbach & Sheuyange BD 3162 (US).