A revision of the Old World Black Nightshades (Morelloid clade of Solanum L., Solanaceae) Author Saerkinen, Tiina Royal Botanic Garden Edinburgh, 20 A Inverleith Row, Edinburgh EH 3 5 LR, United Kingdom tsarkinen@rbge.org.uk Author Poczai, Peter Botany Unit, Finnish Museum of Natural History, University of Helsinki, P. O. Box 7, FI- 00014 Helsinki, Finland Author Barboza, Gloria E. Instituto Multidisciplinario de Biologia Vegetal (CONICET-Universidad Nacional de Cordoba), Casilla de Correo 495, 5000 Cordoba, Argentina Author Weerden, Gerard M. van der Experimental Garden, Radboud University, Faculty of Science Box 49, P. O. Box 9010, 6500 Nijmegen, The Netherlands Author Baden, Maria Max-Planck Odense Center on the Biodemography of Aging and Department of Biology, University of Southern Denmark, Campusvej 55, DK- 5230 Odense M, Denmark Author Knapp, Sandra Department of Life Sciences, Natural History Museum, Cromwell Road, London SW 7 5 BD, United Kingdom s.knapp@nhm.ac.uk text PhytoKeys 2018 2018-07-25 106 1 223 http://dx.doi.org/10.3897/phytokeys.106.21991 journal article http://dx.doi.org/10.3897/phytokeys.106.21991 1314-2003-106-1 FF8BFFC82928FFA84734FFCB2E61E260 1326005 16. Solanum tarderemotum Bitter, Repert. Spec. Nov. Regni Veg. 10: 547. 1912 Figures 49 , 50 Solanum dasytrichum Bitter, Bot. Jahrb. Syst. 49: 568. 1913. Solanum tarderemotum Type. Tanzania. Tanga: Usambara, Kwai, E. Eick 227 (holotype: B, destroyed; no duplicates found). Solanum florulentum Bitter, Repert. Spec. Nov. Regni Veg. 10: 544. 1912. Solanum tarderemotum Type. Tanzania. Tanga: Lushoto, Distr. Kwai, 1600 m, E. Albers 189 (holotype: B, destroyed; no isotype at EA). Solanum kifinikense Bitter, Repert. Spec. Nov. Regni Veg. 10: 545. 1912. Solanum tarderemotum Type. Tanzania. Kilimanjaro: Kifinika volcano, Mar 1894, G. Volkens 1909 (lectotype, designated by Edmonds 2012 , pg. 127: HBG [HBG520843]; isolectotype: BR [BR0000008799418]). Solanum pentagonocalyx Bitter, Repert. Spec. Nov. Regni Veg. 10: 544. 1912. Solanum tarderemotum Type. Tanzania. Tanga: Usambara, Kwa Mstuza, Handei Kewegolot, Aug 1893, C. Holst 9021 (lectotype, designated by Edmonds 2012 , pg. 138: M [M-0105625]; isolectotypes: HBG [HBG520844], W [1894-0006577]). Solanum tetrachondrum Bitter, Bot. Jahrb. Syst. 49: 565. 1913. Solanum tarderemotum Type. Tanzania. Kilimanjaro: near Marangu, G. Volkens 623 (holotype: B, destroyed; no duplicates found). Solanum tetrachondrum Bitter var. subintegrum Bitter, Bot. Jahrb. Syst. 49: 566. 1913. Solanum tarderemotum Type. Tanzania. Kilimanjaro: Marangu, G. Volkens 622 (holotype: B, destroyed; no duplicates found). Solanum viridimaculatum Gilli, Ann. Naturhist. Mus. Wien 77: 43. 1973. Solanum tarderemotum Type. Tanzania. Njombe: Madunda, Livingstone-Gebirge am Nordostufer des Nyassasees, bei Madunda, Nebelwald, 2200 m, 29 Jul 1958, A. Gilli 499 (holotype: W [1973-0001020]). Type . Tanzania . Kilimanjaro : Marangu , 1600 m , 10 Sep 1910 , H.J.P. Winkler 3856 ( holotype : WRSL) . Description. Annual to short lived erect to weakly scrambling perennials to 1.5 m tall, subwoody and somewhat branching at base. Stems spreading to decumbent, usually somewhat winged and with spinescent processes, fleshy, green to somewhat purple tinged, older stems drying pale yellowish-brown or whitish-grey, markedly hollow and even older stems collapsing in herbarium specimens; new growth glabrous or sparsely to moderately pubescent with simple, spreading, uniseriate, translucent, usually eglandular (sometimes tipped with a single-celled gland) trichomes, these 3-5-celled, 0.2-1.0 mm long; older stems glabrous. Sympodial units difoliate, the leaves usually not geminate. Leaves simple, 3-13 cm long, 1.5-2.5 cm wide, ovate to elliptic, very variable in size even on an individual plant, membranous, concolorous, without smell; adaxial surface glabrescent to sparsely pubescent with simple uniseriate trichomes like those on stem; abaxial surface usually less pubescent that the adaxial ones, glabrous to sparsely pubescent with simple uniseriate trichomes; major veins 4-8 pairs; base abruptly attenuate; margins entire or very rarely shallowly sinuate to lobed, if so the tips of the lobes acute or rounded; apex acute to attenuate; petioles (0-)4-8 cm long, pubescent with simple uniseriate trichomes like the leaves. Inflorescences 1-4(-6) cm long, internodal, unbranched or with up to 5 branches, but if branched usually only furcate, the flowers spaced along the rhachis, with 10-40 flowers, glabrous or sparsely pubescent like the stems; peduncle 1-3(14) cm long; pedicels 0.7-0.8 cm long, ca. 0.3 mm in diameter at the base, ca. 0.5 mm in diameter at the apex, slender, nodding at anthesis, sharply bent just above the insertion point so the base of the pedicel is an acute angle, this especially noticeable in fruit, articulated at the base or ca. 1 mm from the rhachis leaving a small stump, falling with the fruit at maturity; pedicel scars 1-2 mm apart in the distal part of the inflorescence, further apart towards the base. Buds ellipsoid, the corolla exserted ca. 1/2 way from the calyx until just before anthesis. Flowers 5-merous, perfect (although style length differs in flowers along the rhachis). Calyx tube ca. 1 mm long, conical with hyaline sinuses, the lobes 0.5-1 mm long, 0.4-0.9 mm wide, ovate to broadly triangular, glabrous to sparsely pubescent like the rest of the inflorescence. Corolla (6-)8-10 mm in diameter, white, stellate, lobed ca. 1/2 way to the base, the lobes 3-4 mm long, 1.5-2 mm wide, spreading or reflexed (apparently reflexed in older flowers), minutely papillate on the tips and margins. Stamens equal; filament tube ca. 0.25 mm long; free portion of the filaments 0.5-0.75 mmm long, adaxially pubescent with tangled simple uniseriate trichomes; anthers 1.5-2.5 mm long, 0.75-1 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary rounded, glabrous; style 3.5-5 mm long, bent or straight, densely pubescent in the basal 1/2 to 1/3, exserted to ca. 1 mm or occasionally included in the anther cone and the stigma flush with the pores; stigma capitate, the surface minutely papillate. Fruit a globose berry, 4-6 mm in diameter, pale green or translucent blackish-grey when ripe, the pericarp thin, matte; fruiting pedicels 0.8-1.1 cm long, ca. 1 mm in diameter at the base, sharply bent at the base and strongly pendent, dropping with mature fruits, not persistent; fruiting calyx not accrescent, the tube ca. 1 mm long, the lobes 0.5-1 mm long, appressed to the berry, the calyx from above pentagonal or stellate. Seeds 20-40 per berry, 1.5-2 mm long, 1-1.5 mm wide, flattened and tear-drop shaped with a subapical hilum, tan or pale brown, the surfaces minutely pitted, the testal cells elongate-rectangular. Stone cells (0-)1-5(-10) per berry. Chromosome number: 2n =4x=48 ( Olet et al. 2015 ; Manoko 2007 ). Figure 49. Solanum tarderemotum A Habit of glabrescent form B Habit of pubescent form C Detail of adaxial leaf surface D Eglandular trichome E Glandular trichome F Inflorescence G Bud H Flower at anthesis I Furcate inflorescence J Infructescence ( A, J Friis et al. 2047 ; B-E Burger 2006 ; F, G Gilbert & Thulin 651 ; H Nijmegen acc. A34750039; I Goode G/72 ). Scale bar: 4 cm ( A-B ), 4 mm ( C ), 0.75 mm ( D-E ), 1 cm ( F, I ), 5 mm ( G-H ) and 1.5 cm ( J ). Drawing by L. Smith. Figure 50. Solanum tarderemotum A Habit B Inflorescence C Buds D Flowers at full anthesis E Infructescence F Mature fruits ( A Nijmegen acc. A54750214; B Nijmegen acc. A14750164; C Nijmegen acc. A34750035; D Nijmegen acc. A34750039; E Nijmegen acc. A54750214; F Nijmegen acc. HNSAR22). Photos by S. Knapp. Distribution (Figure 51 ). Common throughout sub-Saharan tropical Africa from Sudan to Mozambique and South Africa and west to Cameroon and Angola; we have also seen a few isolated collections from the Comoro Islands and Madagascar (perhaps introduced). Figure 51. Distribution of Solanum tarderemotum . Ecology. Grows in a wide variety of disturbed habitats, along forest margins and roadsides and in clearings, often cultivated; between (0-) 500 and 3,300 m elevation. Common names. Burundi: isogo [Kirundi language], urusongo; Cameroon: kifon, kre' fom; Democratic Republic of the Congo: mulunda; Ethiopia: chau, murenchiwa/marenchiwa, nech tunaye; Kenya: abune, isoiik, isoyiot, isusa, litusta/liususa, lokitoemenyan, managu, mnavu, ol' momoye, osuga, sujet, usuga; Rwanda: isogo, ururandayri [Kinyarwanda language]; Tanzania: enyafui [Kimaasai language], mhaki, mnavu, msogo, muhaka, ormomoi aves, soko, suga; Uganda: ensugga, enswiga/eshwiga/eswiga [Kigezi language], nouiga, nsuggaenzirugavu; Zimbabwe: m'sungu sungu-m'adzizambito . Uses. Leaves eaten as a vegetable throughout the range; also used as a medicinal plant (in Karamoja Distr., Uganda). Preliminary conservation status ( IUCN 2016 ). Solanum tarderemotum is widespread across tropical Africa and can be assigned a preliminary status of LC (Least Concern; Table 7 ). Discussion. Solanum tarderemotum is a morphologically variable species. Edmonds (2012) recognised individuals with branched inflorescences as S. florulentum and applied S. tarderemotum only to material with simple inflorescences, but branched and simple inflorescences can occur on the same specimen, depending on age. Intermediate material was occasionally recognised in some accounts under the unpublished name " Solanum eldorettii " ( Olet 2004 ; Manoko 2007 ). Crosses between the forms of S. tarderemotum (as S. tarderemotum A, B, and C, and S. florulentum ) yielded the full range of phenotypes, including intermediate individuals and both "parental" phenotypes ( Manoko 2007 ). Solanum tarderemotum is distinguished from other sympatric species by its usually many-flowered inflorescences, long-exserted style that often is bent rather than perfectly straight (easier to see in live plants), spathulate calyx lobes that are rounded at apex, very strongly reflexed fruiting pedicels and berries that drop with the pedicel attached. Stems of S. tarderemotum are hollow and collapse upon drying, unlike the stems of the similar S. villosum that are more solid. Solanum scabrum also has hollow stems, but fewer-flowered inflorescences and larger fruits that drop without the pedicels. Solanum nigrum (North African populations), S. scabrum and S. villosum all lack stone cells, while the fruits of S. tarderemotum consistently have 2-6 stone cells in each berry. Most material of S. tarderemotum is either glabrous or has eglandular pubescence, but plants from higher elevations in the eastern African mountains are glandular pubescent; these were identified as S. pseudospinosum by Edmonds (2012) . Previous molecular studies identified S. tarderemotum form C (Clade IV) as a separate entity based on AFLP data that was analysed using Maximum Parsimony phylogenetic method ( Manoko 2007 ). Individuals clustering apart from the main clade of S. tarderemotum and S. florulentum (Clade IV) showed simple inflorescences with fewer flowers, narrower corolla lobes and green fragrant fruits at maturity distinct from majority of accessions of S. tarderemotum and S. florulentum ( Manoko 2007 ). The results from the AFLP phylogeny are not, however, supported by sequence based molecular phylogeny based on both rapidly evolving non-coding plastid and low-copy nuclear markers ( Saerkinen et al. unpublished). The accession, on which S. tarderemotum form C (Clade IV) was based (NIJ 964750060), does not have locality or origin data and hence, we prefer assuming that the collection represents morphological variation of S. tarderemotum as circumscribed here, supported by molecular sequence data. The differing relationships in the AFLP phylogeny in Manoko (2007) could be explained by the general difficulty in interpreting AFLP banding patterns objectively in closely related species and highlight the issues in using AFLP data in phylogenetic analyses. Solanum tarderemotum is a tetraploid species with unknown parentage ( Olet et al. 2015 ). The species has not been included in any previous crossing and molecular studies that have focused on understanding the parental origin of the hexaploid species S. nigrum and S. scabrum (e.g. Edmonds 1977 , 1979a ; Ganapathi and Rao 1985 , 1986b , 1986c , 1987a , 1987b ; Jacoby et al. 2003 ; Jacoby and Labuschagne 2006 ; van der Walt et al. 2008 ; Poczai and Hyvoenen 2011 ). It could represent one of the tetraploid parents of the hexaploids and should be included in further studies focused on understanding the origin of the polyploids. Both names S. tarderemotum and S. florulentum were published in the same publication. No publication has yet synonymised the two. Most of the types for all names associated with our circumscription of this species were destroyed in Berlin. We have chosen to use the name S. tarderemotum because the holotype (WRSL) is still extant, while no known duplicates of the type of S. florulentum have been located. No duplicate material has been found for the type collection of S. dasytrichum that was described from a specimen in B that is no longer extant; Edmonds (2012) suggested this was a synonym of S. florulentum with the reservation that the description did not mention forked inflorescences, but otherwise matched. In the protologue, Bitter (1913a) says he originally identified this as S. hildebrandtii A.Braun & C.D. Bouche (= S. villosum ), but that it differs in having 4 stone cells; we suspect this plant is a hybrid between S. villosum and S. tarderemotum , but we place it in synonymy here due to its possession of stone cells (lacking in S. villosum ). Selected specimens examined. Angola . Huila : Ha, juxta ripas rivi de Sopollo , Dec 1859 , Welwitsch 6034 (BM, K) ; Uige : Santa Cruz Mission , 12 Aug 1962 , Codd 7521 (K) . Botswana . North-West : Ngami , Central Management Unit , Selinda Reserve , 8 Apr 2005 , Heath & Heath 1023 (K) ; Burundi . Rwegura , Territoire Kayenza , 28 May 1969 , Lewalle 3620 (K) ; Bugarama : Teza , 29 Dec 1978 , Reekmans 7400 (EA, K, MO) ; Bujumbura : Katumba , 12 Dec 1979 , Reekmans 8444 (MO) ; Muramvya : Nyabigondo , 2 Feb 1967 , Lewalle 1536 (MO) . Cameroon . Nord-Ouest : Bui , Oku-Elak , 27 Oct 1996 , Cheek et al. 8455 (K, MO); Boyo, Ijim Mountain Forest , 21 Nov 1996 , Kamundi , et al. 673 (K, MO) ; Sud-Ouest : Mt Kupe , Nyasoso , 23 Oct 1995 , Cheek et al. 7470 (K, MO); Mt. Cameroon R.C. , 24 Mar 1961 , Swarbrick SCA271 (E) . Cape Verde . Santiago : valley of Santo Domingo , 3 Nov 1839 , Hooker 117 bis (K) . Chad . Lac : Koulfe , Chari Central , 28 Jun 1903 , Chevalier 8788 (K, P); N'Djamena : c. 65 km S of Fort Lamy , 1 Jan 1965 , de Wilde et al. 5118 (K) . Comoros . Iles Comores , Boivin s.n. (W); Mohilla Island , 5 Apr 1861 , Meller s.n. (K); Anjouan : sin. loc, Jun 1875 , Hildebrandt 1626[a ] (BM); Mwali : Insul. Mohely, 1854, Boivin s.n. (BM) . Democratic Republic of the Congo . Katanga : a 3 km de Lukuni (Katanga), source de la Kasapa , 12 May 1961 , Poelman 7 (K); Katuba, ferme Droogmums, Kaletele, Jan 1927 , Quarre 18 (GH) ; Kivu Nord : Ngungu , 18 km SW Sake , 11 Aug 1954 , Stauffer 48 (K, P); Kibati , Nord-Kivu : Butembo , 21 Feb 1974 , Baudet 487 (K); NW slope of Mt. Vislke , 24 Feb 1975 , D'Arcy 8089 (MO) ; Orientale : Ituri , Kibali , Aru , Aug 1931 , Lebrun 3575 (GH, K); Sud Kivu : Lac Tsimuka , plaine de la Ruzizi , Jan 1950 , Germain 5593 (K); Kalonge , Kalonge, long riviere Nyamwamba endroit fran et ombrage, 12 Feb 1953 , de Witte 10477 (MO) . Eritrea . Semienawi Keyih Bahri : c. 10 km S of Nefasit , 2 Feb 1969 , de Wilde 4502 (K) . Ethiopia . Amhara : South Gondar , 4 km N of Debre Tabor , 13 Sep 2004 , Friis et al. 11552 (K); Semien Mountains , 1 Oct 2003 , Wieringa 4971 (K) ; Dire Dawa : Dire Dawa , 16 Oct 1969 , Parker 580 (K) ; Harari : between Harrar and Abbaba , Sep 1901 , Wellby s.n. (K) ; Oromia : Gara Mullata , c. 50 km due W of Harar , 2 Aug 1962 , Burger 2006 (FT, K); Hana , E part of the Omo , 23 Mar 1976 , Fukui 15 (EA, K); c. 5 km N of Addis Ababa , 3 May 1965 , de Wilde & de Wilde-Duyfjes 6507 (K, MO) ; Somali : Harerge , on the rd from Alemaya to Asbe Tafari , 6 km from Kobbo , 16 Aug 1967 , Westphal & Westphal-Stevels 1227 (K) ; Southern Nations (SNNP) : Lower Omo River , 7 Oct 1970 , Carr 875 (EA, K); rd from Jimma to Serbo, 14 km from Jimma, 1 Aug 1968 , Westphal & Westphal-Stevels 5499 (K, MO) . Ghana . Ohamu , Agric. Res. Station , Jul 1961 , Irvine 4957 (K) . Guinea . Nzerekore : Nzo , Mt Nimba , 5 Nov 1969 , Adam 24671 (MO) . Kenya . Central : vicinity of Lake Naivasha , 17 Jul 1909 , Mearns 842 (GH); Meru North , Mt. Kenya area , 18 Apr 2010 , Vorontsova et al. 198 (BM, BR, EA, K, MO, NY) ; Coast : Mwanda , Mgange Nyika , 3 Oct 1971 , Klungness 74 (K) ; Eastern : Makueni , Chyulu Hills , 15 May 1938 , Bally 7787 (EA, K) ; Nairobi : Nairobi , 1924, McDonald 808 (K); nr Nairobi , Whyte s.n. (K) ; Rift Valley : Mt. Elgon , Kitale , 27 Dec 1960 , Loeffler E-107 (W); Kericho , Southwestern Mau Forest Reserve , camp 7, river Dimbilil , 7 Aug 1949 , Maas Geesternanus 5603 (K, MO); Katilia forest , 12 mi NNE of Kangetet , Kerio River, 25 May 1970 , Mathew 6379 (EA, K) ; Western : Kakamega , Kaptiki Secondary School , 11 Nov 1984 , Hohl 157 (EA, W); pr. Forest Station ad mar, Mt Aberdare Expedition , 14 Jan 1922 , Rob & Fries 921 (MO) . Lesotho . Sehlabathebe , 4 Jan 1973 , Guillarmod et al. 138 (K) . Madagascar . Antananarivo : Ambohidratrimo, Ambohimanga, pres de Tananarive , 19 Apr 1928 , Decary 6179 (P); Antananarivo-Nord, Tananarive, Waterlot s.n. (P) ; Toliara : valley half a mi W of Ampoza , 5 Sep 1929 , White s.n. (BM) . Malawi . Central : Dedza Distr. , Dedza mountain forest, 19 Jan 1987 , Balaka & Patel 1861 (MO) ; Northern : Rumphi Distr. , Nykia Plateau , 2 mi E of Chelinda , 4 Mar 1977 , Pawek 12428 (K, MO) ; Southern : Mt. Mulanje , foot NE slopes of Namasile opposite Sombani hut, 3 Jan 1971 , Hilliard & Burtt 6134 (E) . Mali . Ackerrand kurz vor Koulikoro , 29 Sep 1992 , Ehrich 336 (B) . Mozambique . Maputo : Lourengo Marques , Namaacha , 1 Aug 1967 , Marques 2142 (MO) ; Zambezia : Namuli Mountain , Muretha Plateau , 26 May 2007 , Harris 186 (K) . Namibia . Kavango West : Tondoro Mission , Tondoro Camp 1 km E of Mission , 15 Dec 1955 , de Winter 3955 (K) ; Zambezi : E Caprivi , Zipfel , Lizazuli , 2 Jan 1959 , Killick & Leistner 3253 (K) . Nigeria . Bauchi : Toro , Panshanu Pass , 15 Aug 1962 , Lawlor & Hall 410 (K) ; Cross River : probably collected nr Obudu Cattle Range , 25 May 1971 , Meer van , 1861 (MO) ; Enugu : Nsukka , University of Nigeria campus, 23 Jan 1962 , Okigbo , 57 (K) ; Osun : Shasha Forest Reserve , 23 Apr 1968 , Gledhill , 998 (K) . Rwanda . Northern : Mt. Visoke , 6 Apr 1970 , Fossey 16 (EA, K) . Senegal . Nr. Bono , nec non Ins. Bonavista, Brunner s.n. (K); Dec 1823 , Roger 17 (K) . Sierra Leone . Northern : Mt. Bintumani , Kabala (admin), Mt Loma , Mira , 30 Nov 1965 , Adam 22258 (GH, MO) . Somalia . Banaadir : 3 km from Muqdisho airport along rd to Jasiira , 4 May 1990 , Thulin & Hedren 7172 (K) ; Togdheer : Wagga Mt. , 1905, Bury s.n. (BM) ; Woqooyi Galbeed : Murak , Sep 1933 , Godding 162 (K); mountains above Qoton, 11 Feb 2002 , Thulin 10906 (K) . South Africa . Eastern Cape : Somerset , 1860, Cooper 528 (K); Fort Beaufort , 1860, Cooper 554 (K, W); Uitenhage , Enon , Drege s.n. (K) ; Gauteng : Pretoria , Region SWA, Kaokoveld , Otjomborombonga on Kunene River , 13 Jul 1976 , Leistner et al. 104 (K, MO) ; KwaZulu-Natal : Weza , Ingeli , 5 Mar 1972 , Strey 10899 (K, MO) ; Mpumalanga : Matebe Valley , May 1883 , Holub s.n. (K) ; North West : nr Vryburg , 60 mi NW of Vryburg , Kalahari Desert , 5 Feb 1948 , Rodin 3500 (K, MO) . South Sudan . Bahr El Ghazal : Isablei [?], Lande der Bongo , 26 Nov 1869 , Schweinfurth 2649 (K) ; Equatoria : Imatong Mountains , Mt. Angargi , 14 Jun 1939 , Andrews 1947 (K); Distr. Torit, Lowiliwili, Imatongs, 14 Nov 1949 , Jackson 902 (BM); Greater Upper Nile : Upper Nile , Boing , 26 Oct 1951 , Sherif A 2891 (K) . Sudan . Blue Nile : Sennar , Wuertemmburg s.n. (W); Darfur : Jebel Marra , Nyertete , 21 Jan 1964 , Wickens 1044 (K); Jebel Marra, Zalingei, Wickens 1776 (K) ; Kassala : Erkowit , Red Sea Hills , Mar 1929 , Lady Maffey 39 (K); Nr Kamobsana, Red Sea Prov. , 25 Jan 1912 , MacDougal & Sykes 137 (BM); Kamobsana, Red Sea Prov. , 25 Jan 1912 , MacDougal & Sykes 139a (BM). Kurdufan : ad Cordofanum Milbes, 4 Dec 1839 , Kotschy 291 (BM, E, GH, K, P, W) . Tanzania . Arusha : Mbulmbul , Block DL , 49, 24 Jun 1944 , Greenway 6951 (EA, K); Ngorogoro Crater, 24 Jun 1938 , Pole Evans & Erens 936 A (E, K, P); Buha : Kakombe Valley , E shore of Tanganyika Lake from Gombe stream to Missonge, 25 Dec 1963 , Pirozynski P 86 (EA, K); Eastern : Morogoro , Lukwangulu Plateau , Uluguru Mts , 19 Sep 1970 , Thulin & Mhoro 1048 (K) ; Iringa : Kidatu , Iringa District , E dam site, 28 Mar 1971 , Mhoro 864 (EA, K) ; Kagera : Karagwe , 1 Mar 1862 , Speke & Grant 453 (K) ; Kigoma : Mpanda , Sisaga , Mahali Mountains , 27 Aug 1958 , Jefford et al. 1818 (K); Kakombe , 10 mi W of Kigoma , 7 Jul 1959 , Newbould & Harley 4285 (K, MO) ; Kilimanjaro :, Chome Forest Reserve , Namboja , 30 Mar 2001 , Mlangwa et al. 1506 (MO) ; Mbeya : Mbeya , 3 Mar 1932 , Davies 461 (EA, K); Rungwe , Nyassa Hochland-Station Kyimbila , 1911, Stolz 384 (B, BM, K, MO, W) ; Mbeya / Njombe : Poroto Mountains , Mbeya Distr. , 16 May 1957 , Richards 9735 (EA, K); Mbulu : Lake Manyara National Park , 2 Dec 1963 , Greenway & Kirrika 11113 (EA, K); Hanang Mt. , 3 May 1962 , Polhill & Paulo 2302 (EA, K) ; Morogoro : Kiberege , Mar 1936 , Culwick 2 (K); Kanga Mountains , 11 May 2007 , Luke et al. 12026 (EA, K, MO) ; Mwanza : Mwanza , Davis 201 (K); Njombe : Stromgebeit des obern Ruhudje , Landschaft Lupembe , nordlich des Flusses , Mar 1931 , Schlieben 413 (K); Northern : Moshi, Weru-Weru gorge, 22 Feb 1955 , Huxley 116 (EA, K) ; Rukwa : Nsanga Forest , Mpanda , Ufipa , 8 Aug 1960 , Richards 13004 (K) ; Ruvuma : Songea , Matagoro Hills , just S of Songea , 22 Feb 1956 , Milne-Redhead & Taylor 8869 (EA, K) ; Shinyanga : Shinyanga , Nov 1938 , Koritschoner 1903 (EA, K) ; Tabora : Unyamwezi , Mininga , 1860, Speke & Grant 79 (K) ; Tanga : Usambaras , between Ngua and Magunga Estates , 17 Jul 1953 , Drummond & Hemsley 3347 (EA, K); Ufipa : Sumbawanga, 3 km SE of Moravian Mission at Nkutwe nr Tatanda, 31 Oct 1992 , Harder & Kayombo 1351 (EA, MO); Ulanga : Ifakara, 16 Jul 1959 , Haerdi 286o (EA, K) . Uganda . Central : Mukono , Kipayo , Dec 1913 , Duemmer 563 (BM, K, P); Mengo , 12 mi to Kampala , Entebbe rd, May 1932 , Eggeling 697 (EA, K) ; Eastern : Mt. Elgon , 25 Jan 1993 , Katende & Sheil 1099 (K) ; Northern : Karamoja , Moroto Township , Sep 1958 , Wilson 609 (EA, K) ; Western : Kigezi D.F.I , 28 Aug 1972 , Goode G-2-72 (K); Mihunga, Ruwenzori, 12 Jan 1939 , Loveridge 350 (A, K, MO); Kigezi, Muhavura Hill, 11 Jan 1933 , Rogers 337 (BM, EA, K) . Zambia . Lusaka : Mt. Makulu , Kafue Basin , 10 Apr 1963 , van Rensberg 1886 (K) ; North-Western : Mwinilunga , 4 Oct 1937 , Milne-Redhead 2565 (K) ; Northern : below Kwimbi Mission , 10 Feb 1955 , Richards 4426 (K) ; Southern : Livingstone Distr. , Victoria Falls , 21 Feb 1997 , Luwiika et al. 460 (MO) . Zimbabwe . Harare : Salisbury , 26 Feb 1927 , Eyles 4712 (K) ; Manicaland : Nr Chirinda , May 1906 , Swynnerton 481 (BM, K) ; Mashonaland Central : Mazowe , Umvukwes , 17 Dec 1952 , Wild 40771 (K) ; Mashonaland East : Distr. Salisbury , Mandara , 25 Sep 1974 , Bisgel 4632 (MO); Marandella , 8 Apr 1948 , Corby 1-20917 (K) ; Matabeleland South : Mazowe , Matobo National Park , Matopus NP, 24 Feb 1981 , Philcox & Leppard 8823 (K) .