A taxonomic revision of Stenoglottis (Orchideae, Orchidoideae, Orchidaceae) Author Phillips, Dean P. 0000-0002-8355-5081 Bews Herbarium, Centre for Functional Biodiversity, School of Life Science, University of KwaZulu-Natal, Private Bag X 01, Scottsville 3209, South Africa. & Current address: Bolus Herbarium, Department of Biological Sciences, University of Cape Town, 7701 Rondebosch, South Africa. & dean. phillips @ uct. ac. za; https: // orcid. org / 0000 - 0002 - 8355 - 5081 dean.phillips@uct.ac.za Author Bytebier, Benny 0000-0002-4661-5727 Bews Herbarium, Centre for Functional Biodiversity, School of Life Science, University of KwaZulu-Natal, Private Bag X 01, Scottsville 3209, South Africa. & bytebier @ ukzn. ac. za; https: // orcid. org / 0000 - 0002 - 4661 - 5727 bytebier@ukzn.ac.za text Phytotaxa 2020 2020-08-25 456 3 219 243 http://dx.doi.org/10.11646/phytotaxa.456.3.1 journal article 3961 10.11646/phytotaxa.456.3.1 e20afa6a-b9b0-40d6-922a-349da64e1a5b 1179-3163 5585921 1.1. Stenoglottis fimbriata Lindl. var. fimbriata ( Figs 1–3 ) Heterotypic synonyms: Stenoglottis fimbriata subsp. saxicola (Schltr. ex Kraenzl.) McDonald (2008: 11) . Stenoglottis fimbriata var. saxicola Schlechter ex Kraenzlin (1898: 567) . Type:— SOUTH AFRICA . KwaZulu-Natal : Mount West, 1585 m , 20 Feb 1895 , Schlechter 6820 ( holotype : B, destroyed; lectotype : BOL, 96151!, here designated; isolectotypes: GRA, 0005004!, NH, 0013312-0!, NY, 00468316 photo!, P, 00339731 photo!, PRE, 0036838-0!, PRE, 0856961-0!). Stenoglottis zambesiaca Rolfe (1897: 190) . Type :— MALAWI . Shire Highlands , 1891, Buchanan 385 ( holotype : K, 000415557!; isotype: BM, 000034610!). Diagnostic characters:— Varieties of S. fimbriata can be distinguished by a combination of their leaf and labellum morphology. The leaves of S. fimbriata var. fimbriata are undulate and variably spotted (from completely unspotted to heavily spotted with brown or maroon; Fig. 1O–P ), whereas in S. fimbriata var. modesta they are always entire, never spotted and somewhat broader and fleshier than those of the nominal variety. In S. fimbriata var. fimbriata , the side lobes of the labellum are also highly variable (in width and degree of fimbriation; Fig. 1A–H, J, L–M ), but in S. fimbriata var. modesta they are always narrow and entire. Flowers of S. fimbriata var. fimbriata are completely spurless, the base of the labellum being consistently flat and smooth ( Fig. 1I, K ), whereas in S. fimbriata var. modesta the tissue at the base of the labellum is slightly protruding, forming what appears to be a minute vestigial spur. Finally, the lifespan of individual flowers of S. fimbriata var. fimbriata is variable (usually 2–3 days, but up to two weeks), whereas in S. fimbriata var. modesta the flowers seldom remain open for more than two days. Stenoglottis fimbriata var. fimbriata is easily distinguished from S. inandensis , S. woodii , and S. macloughlinii by its spurless labellum and undulate leaves. It is most likely to be confused with S. longifolia but differs in the orientation of its floral bracts and various characteristics of the column. For a detailed diagnosis of S. fimbriata (both varieties) and S. longifolia , see ‘diagnostic characters’ under S. longifolia . FIGURE 1 . Selected specimens of Stenoglottis fimbriata var. fimbriata .A, inflorescence, Ngome Forest Reserve (KZN). B–C, inflorescence and flower ventral view, Umtamvuna Nature Reserve (KZN). D, flower ventral view, Msikaba (EC). E, flower ventral view, Ingomankulu (KZN). F, flower ventral view, Highflats (KZN). G, flower ventral view, Lekgalameetse Nature Reserve (LIM). H, flowers, Mount West (KZN). I–J, flower lateral view (showing spurless labellum and sheathing bract) and flower ventral view, Kaapschehoop (MP). K–L, flower lateral and ventral view, Winston Park (KZN). M, flowers, Signal Hill, Grahamstown (EC). N, habitat, Royal Natal National Park (KZN). O, leaf rosette ( in situ ), Kei Mouth (EC). P, leaf rosette ( in situ ), God’s Window (MP). Scale bars = 5 mm. N–P reproduced with the permission of Herbert Stärker. FIGURE 2. Selected specimens of Stenoglottis fimbriata var. fimbriata . A, column lateral view, Oribi Gorge Nature Reserve (KZN). B–C, column lateral and ventral view, Msikaba (EC). D, column lateral view, Nkandla Forest Reserve (KZN). E–F, column lateral and ventral view, Kaapschehoop (MP). Scale bars = 0.5 mm. Abbreviations: au = auricle, an = anther, stg = stigmatic arm. Notes:— Stenoglottis fimbriata is a widespread and morphologically variable taxon, and previous attempts to subdivide it are largely unsupported by morphological or phylogenetic evidence. Stenoglottis fimbriata var. saxicola was not recognised by Stewart (1989) but was by McDonald (2008) , who transferred it to the rank of subspecies. This taxon (as both variety and subspecies) was regarded as a diminutive, highland form. Plants collected from higher elevations are indeed often smaller than lowland plants, and this size difference is to some extent maintained in cultivation and therefore mostly genetic ( Stewart 1989 , this study). However, changes in overall plant size with elevation are gradual, and consequently their separation on this basis is arbitrary. In addition to plant size, McDonald (2008) attributed several other morphological differences to S. fimbriata subsp. saxicola : colony-forming habit, stigmas are shorter than the anthers, narrower median sepals and more falcate petals. However, these characters too have been shown to be variable and inconsistent as diagnostic characters. Multivariate morphometric analyses did not resolve the subspecies of S. fimbriata as discrete morphological clusters, and, likewise, there is no phylogenetic basis to support their distinction ( Phillips & Bytebier 2020 ). Stenoglottis fimbriata subsp. saxicola is therefore not recognized here. The distinction of S. zambesiaca from S. fimbriata is also problematic for several reasons. Its original description by Rolfe (1897) vaguely stated that it differs from S. fimbriata “in the shape of the leaves, and in various other details”. It was later synonymized with S. fimbriata by Summerhayes (1968) and then resurrected by Stewart (1989) , who expanded on Rolfe’s diagnosis, distinguishing it from S. fimbriata on the basis of its smaller flowers with fewer, larger spots, a dorsal sepal that is much narrower than the laterals, labellum side lobes that are ligulate and round or truncate apically and auricles that are tuberculate at the club-shaped apex (as opposed to the slender auricles of S. fimbriata ) . Later, McDonald (1995 , 2006 , 2008 ) attributed to S. zambesiaca the additional characteristics of having markedly falcate petals, a labellum with a narrow claw, and non self-pollinating flowers due to the presence of auricles (sometimes referred to instead as staminodes), which he claimed to be completely absent in S. fimbriata . However, the holotype of S. zambesiaca ( Buchanan 385 ) clearly does not exhibit all qualities that were later ascribed to it. Specifically, the side lobes of its labellum are not particularly round or truncate, and the specimen exhibits a high proportion of seed-set (i.e. many large, swollen capsules) and was therefore almost certainly self-pollinating. The latter was noted by McDonald (1995 , 2008 ), who remarked that even if the plants currently referred to S. zambesiaca do in fact represent a taxonomic entity distinct from S. fimbriata , the name itself is still misapplied. In a published study, S. zambesiaca could neither be distinguished phylogenetically from subspecies of S. fimbriata nor resolved as a discrete group in the morphometric analyses ( Phillips & Bytebier 2020 ). The supposed differences in floral macro-morphology described by Stewart (1989) and McDonald (1995 , 2006 , 2008 ) all fall within the range of variation observed for S. fimbriata . Regarding the auricles, ~300 flowers from specimens of the S. fimbriata complex (i.e. S. modesta , S. zambesiaca and both subspecies of S. fimbriata , as designated in Phillips & Bytebier 2020 ) were dissected and examined. The auricles were found to be rarely absent in individual flowers (and occasionally on one side of the column only) and never completely absent within a population. In these plants, the auricles are usually approximately uniform in width or widen slightly toward the apex but are never as obviously swollen at the apex as they are in the rest of the genus. They also clasp the sides of the anther and therefore do not obstruct the stigmatic arms, which rise up to touch the pollinia. As a result, these flowers are self-pollinating, including those identified as S. zambesiaca (as defined by Stewart and McDonald , e.g. Phillips 29 from Ngome Forest), although the process does seem to be much delayed in some specimens and the proportion of self-pollinating flowers is variable. Presently, there is little published research on pollination in Stenoglottis ( Schelpe 1970 , 1985 ), and this aspect of its biology warrants further study. However, it does not appear that S. fimbriata could be reasonably subdivided on this basis. Given that the type specimen of S. zambesiaca falls within the range of morphological variation of S. fimbriata and that there is no phylogenetic basis for their distinction, S. zambesiaca is also considered here a synonym of S. fimbriata var. fimbriata . FIGURE 3. Distribution of Stenoglottis fimbriata var. fimbriata . Distribution and ecology:— Stenoglottis fimbriata var. fimbriata is widespread throughout southern and east Africa, occurring in South Africa , Eswatini ( Swaziland ), Zimbabwe , Mozambique , Malawi and the southern highlands of Tanzania ( Fig. 3 ) at elevations of 20–2200 m . In South Africa , it occurs in the Eastern Cape (EC), KwaZulu-Natal (KZN), Limpopo (LIM), and Mpumalanga (MP) provinces. There has also been an outlier population recorded from Gabon (not mapped). It occurs in forest or on forest margins in deep or partial shade, most commonly as a lithophyte on mossy boulders or on ledges and crevices along south-facing cliff faces; occasionally it is an epiphyte on tree trunks or terrestrial in shallow soil. It flowers January–April and exhibits a high rate of self-pollination. Specimens:— ESWATINI . Hhohho Region : 2631 ( Mbabane ): Forbes Reef , 1524 m (– AA ) , 7 Apr 1965 , Compton 32331 ( NBG0082066 -0!); Black Mbuluzi Valley , The Caves , 1219 m (– AC ) , 17 Mar 1956 , Compton 25800 ( NBG0057172 -0!, NH0111186 -0!); Hill NE of Mbabane , 1524 m (– AC ) , 2 Mar 1959 , Compton 28549 ( K001208125 !, NH0111185 -0!); Hill NW of Mbabane , 1371 m (– AC ) , 17 Feb 1961 , Dlamini s.n. ( NBG0049288 -0!); Kampaula Valley (– AC ) , Apr 1917 , Edwards 14319 ( BOL 50007!) . GABON . Ngounié Province : Mont Iboundji , on summit, ± 10 km on track to Moughoungoulou Falls , 850 m , 16 Feb 1988 , Louis 2805 ( MO3853478 !) . MALAWI . Central Region : Dedza District , Dedza Mountain , 2140 m , 20 Mar 1955 , Exell et al. 1077 ( BM000034601 !) ; Dedza District , Dedza Mountain , near radio relay station, 2000 m , 3 Mar 1977 , Grosvenor & Renz 1005 ( BOL 49890!, K001208141 !) ; Dedza District , Dedza Mountain , near summit, 2050 m , 5 Apr 1978 , Pawek 14281a ( MA , MO !) . Northern Region : Mzimba District , Vipyha Plateau , Kawandama , 3 Apr 1981 , Sallibeni 2961 ( MO !) ; Rumphi District , Nyika Plateau , Kasyaula Forest , 2000 m , 22 Feb 1982 , Dowsett-Lemaire 354 ( K001208146 !) . Southern Region : Blantyre District , Ndirande , 3 Mar 1961 , Morris 113 ( K001208138 !) ; Chiradzulu District , Chiradzulu Mountain , above Lisao Forest , 1400 m , 13 Mar 1977 , Brummitt et al. 14849 ( K001208140 !) ; Chiradzulu District , Chiradzulu Mountain , 1450 m , 6 Apr 1985 , La Croix et al. 690 ( K001208147 !) ; Mulanje District , Mount Mulanje [ Mlanje ], 1897, Adamson 433 ( K001208143 !, P00339730 photo!) ; Mulanje District , Tuchila Plateau , May 1901 , Purves 5 ( K001208142 !) ; Zomba District , Malosa Crater , 762 m , Morris 93 ( K001208139 !) ; Zomba District , Zomba Mountain , 1550 m , 31 Mar 1978 , Pawek 14170 ( K , MA , MO !, SRGH , UC ) ; Zomba District , Zomba Mountain , 26 Feb 1981 , Welsh 669 ( K001208145 !) ; Zomba District , Zomba Mountain , Westwood 620 ( K001208144 !) . Without precise locality : Southern Region , Shire Highlands , 1881 Buchanan s.n. ( K000415558 !) . MOZAMBIQUE . Sofala Province : 1834 ( Vila Paiva de Andrada ) : Gorongosa District , Mount Gorongosa, SE slopes, 1400 m (– AC ) 23 Jul 1970 , Müller & Gordon 1428 ( K001208154 !) ; Gorongosa District , Mount Gorongosa , 1768 m (– AC ) , Schelpe s.n. ( BOL 49906!) . Zambezia Province : 1536 ( Gurué ) , Gurué District , Serra de Gurué , 1280 m (– BD ) , 24 Jul 1979 , De Koning 7430 ( BM000034602 !, BR000002181789 photo!, MO!, WAG1140769 photo!) . SOUTH AFRICA . Eastern Cape : 3029 ( Kokstad ): Mount Ayliff , Insizwa Forest (– CC ) , 22 Feb 1972 , Strey 10717 ( PRE0111899 -0!); Mount Ayliff , Insizwa Mountain (– CD ) , 17 Feb 1971 , Hilliard & Burtt 6570 ( E , K , MO !, NU0010087 !) . 3127 ( Lady Frere ): Engcobo , 1275 m (– DB ) , 15 Feb 1912 , McLoughlin 47b ( BOL 49893!) . 3128 ( Umtata ): Maclear , Rockwater Farm, NE mountain slope, 1550 m (– AA ) , 23 Feb 1993 , Bester 164 ( PRE0804517 -0!); Maclear , Farm 164, ± 17 km north of Ugie , 1420 m (– AA ) , 23 Feb 1994 , Bester 2323 ( NH0118861 -0!); Maclear , Farm 164, ± 17 km north of Ugie , 1420 m (– AA ) , 23 Feb 1994 , Bester 2324 ( PRU !) ; Mnyolo , on road between Mnyolo and Ugie , 993 m (– AD ) , 15 Jan 2017 (ex hort. Feb 2018 ), Phillips 51 ( NU0088388 !) ; Maclear , between Tsolo and Maclear (– BA ) , 2 Mar 1979 , Linder 2103 ( BOL 49921!) ; Mhlahlane Forest Station , Rubuxa Mountain , NW of Umtata , 1600 m (– BC ) , 27 Feb 1997 , Dold & Cocks 2806 ( GRA0005019 !) . 3129 ( Port St Johns ): Msikaba , on bank of Msikaba River near mouth, 21 m (– BD ) , 16 Dec 2016 (ex hort. Apr 2018 ), Phillips & Kiepiel 46 ( BNRH !, NU0088383 !) ; Port St Johns (– DA ) , May 1911 , Mason s.n. ( K001208137 !) ; Port St Johns (– DA ) , 1900, Pegler 542 ( PRE0036835 -0!); Port St Johns (– DA ) , 9 Mar 1950 , Prance NBG339/38 ( NBG0057173 -0!); Port St Johns (– DA ) , 8 Apr 1938 , Prance s.n. ( NBG0057176 -0!); Port St Johns , near township (– DA ) , Dec 1943 , Schelpe 315 ( NU0010067 !) . 3130 ( Port Edward ): Mtentu River gorge, north bank (– AA ) , Jun 2006 , Edwards 3271 ( NU0016759 !) . 3226 ( Fort Beaufort ): Katberg (– BC ) , Mar 1885 , Baur 1105 ( K001208135 !) ; Katberg (– BC ) , date unknown, Hutton s.n. ( K001208136 !) ; Fort Fordyce , 920 m (– CB ) , 7Apr 2015 , Bodeux 21 ( GRA0005026 !) ; Fort Fordyce , 1157 m (– CB ) , 21 Mar 2014 , Martin 5 ( GRA0005025 !) ; Fort Fordyce Nature Reserve , 1041 m (– CB ) , 14 Jan 2017 (ex hort. Mar 2018 ), Phillips 49 ( NU0088386 !) ; Fort Fordyce , 914 m (– CB ) , 12 Mar 1947 , Story 2111 ( PRE0036834 -0!). 3227 ( Stutterheim ): Hogsback , Boma Pass (– CA ) , 3 Apr 1943 , Acocks 9744 ( PRE0036828 -0!); Hogsback , Swallowtail Falls , 1219 m (– CA ) , 10 Mar 1968 , Ashton 194 ( GRA0005014 !) ; Hogsback , Katberg Pass , 1371 m (– CA ) , 3 Apr 1965 , Bokelmann s.n. ( BOL 49904!) ; Stutterheim , Leopard Falls , 1009 m (– CB ) , 19 Jan 2002 , Bytebier et al. 2198 ( NBG0196828 -0!); King William’s Town , Pirie Forest (– CC ) , Mar 1910 , Godfrey 72 ( GRA0005001 !) ; King William’s Town , Pirie Forest (– CC ) , Feb 1944 , R.U.C. Biol. Exped. RUH528 ( GRA0005009 !) ; King William’s Town , Pirie Forest (– CC ) , 9 Mar 1950 , Werner NBG100/49 ( NBG0057174 -0!); King William’s Town , Pirie Forest (– CC ) , 14 Feb 1944 , Widdicombe 8 ( BOL 49900!) ; Yellowwood Falls , 365 m (– CD ) , 1893, Sim 799 ( NU0010058 !) ; Komga , 609 m (– DB ) , Mar 1891 , Flanagan 644 ( GRA0005003 !, PRE0036832 -0!, SAM0020465 !); Komga , 182 m (– DB ) , Mar 1894 , Flanagan 2313 ( PRE0036827 -0!). 3228 ( Butterworth ): Elliotdale , The Haven (– BB ) , 4 Apr 1966 , Gordon-Gray 206 ( NU0010085 !) ; Elliotdale , The Haven (– BB ) , 5 Apr 1966 , Gordon-Gray 242 ( NU0010081 !) ; Kentani , 457 m (– CB ) , Apr 1902 , Pegler 275a ( BOL 96150!, PRE0111898 -0!,); Kentani , 365 m (– CB ) , Apr 1902 , Pegler 275b ( PRE0036829 -0!); Kentani (– CB ) , Mar 1902 , Pegler s.n. ( BOL !) . 3326 ( Grahamstown ): Rabbits Wood , 10 miles from Grahamstown on East London road, 609 m (– BC ) , 12 Mar 1968 , Ashton 193 ( GRA0005016 !) ; Grahamstown , Mountain Drive , 736 m (– BC ) , 19 Mar 1994 , Dold 925 ( GRA0005018 !) ; Grahamstown , Signal Hill (– BC ) , Apr 1896 , Farquhar s.n. ( GRA0005005 !) ; Grahamstown , Signal Hill , 731 m (– BC ) , 1888, Galpin 56 ( BOL 50015!) ; Grahamstown , 670 m (– BC ) , MacOwan 515 ( NY04170350 photo!, SAM0020464-1 !, SAM0020464-2 !); Grahamstown , Signal Hill , 710 m (– BC ) , 10 Jan 2017 (ex hort. Mar 2018 ), Phillips 48 ( BNRH !, NU0088385 !) ; Grahamstown , Featherstone Kloof , 600 m (– BC ) , Feb 1984 , Weeks 134 ( GRA0005017 !) ; Grahamstown (– BC ) , 13 Apr 1944 , Widdicombe 9 ( BOL 49899!) ; Alexandria Forest (– CB ) , May 1956 , Collett s.n. ( GRA0005012 !) . 3327 ( Peddie ): East London (– BB ) , Pearson 1549 ( SAM0001105 -0!). KwaZulu- Natal : 2730 ( Vryheid ): Pongola Bush Nature Reserve , 1700 m (– DB ) , Feb 1993 , Scott-Shaw & Wallace 5187 ( CPF0021094 !) ; Vryheid Hill Nature Reserve , 1302 m (– DD ) , 17 Mar 2016 (ex hort. Mar 2018 ), Phillips 28b ( NU0088334 !) . 2731 ( Louwsberg ): Ngoje Plateau , summit area (– CB ) , 20 Feb 1991 , Scott-Shaw s.n. ( CPF0021103 !) ; Ngome , above forest near Police Post , 1295 m (– CD ) , 3 Apr 1977 , Hilliard & Burtt 9999 ( NU0010086 !) ; Ngome Forest Reserve , 1134 m (– CD ) , 17 Mar 2016 (ex hort. Mar 2018 ), Phillips 29 ( NU0088335 !) ; Ngome Forest Reserve [Ngome State Forest] (– CD ) , 17Apr 2005 , Styles 2612 ( NU0016078 !) . 2828 ( Bethlehem ): Mont-Aux-Sources, Pastures Bush , 1829 m (– DB ) , 16 Feb 1926 , Bayer & McClean 116 ( PRE0036847 -0!); Mont-Aux-Sources, Tugela Valley (– DB ) , Apr 1934 , Humbert 14847 ( P00339724 photo!, P00339725 photo!); Royal Natal National Park , about 1 km north of the Mahai Campsite and Caravan Park , Gudu Bush , 1700 m (– DB ) , 23 Feb 1992 , Kurzweil 1621 ( NBG0150820 -0!); Mont-Aux-Sources, forest patch upstream from hotel (– DB ) , 22 Feb 1979 , Linder 2079 ( BOL 49922!) ; Royal Natal National Park , Tugela Gorge Trail , 1757 m (– DB ) , 21 Feb 2016 (ex hort. Mar 2018 ), Phillips 20 ( NU0088327 !) ; Royal Natal National Park , Tugela Gorge Trail , 1757 m (– DB ) , 21 Feb 2016 (ex hort. Mar 2018 ), Phillips 21 ( NU0088328 !) ; Royal Natal National Park , Tendele Forest (– DB ) , 21 Feb 1976 , Physick 16 ( NU0010088 !) ; Mont-Aux-Sources, Royal Natal National Park (– DB ) , Feb 1964 , Trauseld NU20 ( NU0010083 !) . 2829 ( Harrismith ): Oliviershoek , 1524 m (– CA ) , Feb 1891 , Thode 8160 ( NH0106447 -0!); Cathedral Peak Hotel , 1570 m (– CC ) , 9 Feb 1982 , Bamps 7217 ( BR0000021817830 photo!); Cathedral area (– CC ) , Jan 1941 , Burchell 50 ( NU0010066 !) ; Cathedral Peak area , Nkohlonhla Valley , 1524 m (– CC ) , Jan 1944 , Bursell s.n. ( NU0010063 !) ; Cathedral Peak , 3 km down Mike’s Pass, NW facing slope near Ndumeni River bank (– CC ) , 11 Feb 1984 , Buthelezi 359 ( NH0083444 -0!); Cathedral Peak area , 1524 m (– CC ) , 6 Feb 1946 , Cross 62 ( NU0010060 !) ; Cathedral Peak , Mazonjwana River Valley , 1722 m (– CC ) , 6 Feb 1951 , Killick 1394 ( CPF0021097 !) ; Cathedral Peak , 1480 m (– CC ) , 10 Feb 1982 , Lambinon & Reekmans 82/456 ( BR0000021817854 photo!, PRE0618335 -0!); Cathedral Peak area , Nkohlonhla River , 1493 m (– CC ) , Feb 1943 , Schelpe 191 ( NU0010070 !) ; Cathedral Peak , Ndumeni Forest [Indumeni], 1829 m (– CC ) , 6 Feb 1957 , Schelpe 6348 ( BOL 49918!) ; Cathedral Peak , Ndumeni Forest [ Indumeni ] (– CC ) 27 Jan 1963 , Vermeulen s.n. ( L1526715 photo!). 2830 ( Dundee ): Qudeni Forest Reserve (– DB ) , Feb 1940 , Fisher & Schweickerdt 103 ( NU0010061 !) ; Qudeni , Ekombe Forest , 1524 m (– DB ) , 12 Feb 1945 , Fisher 785 ( NU0010064 !) ; Qudeni , Ekombe Forest , 1524 m (– DB ) , 15 Feb 1945 , Fisher 822 ( NU0010065 !) ; Kranskop , 42 km south of Silutshana on road to Kranskop , 1430 m (– DD ) , 23 Feb 1974 , Davidse 6902 ( MO2413230 !) . 2831 ( Nkandla ): Nkandla Forest Reserve , 1197 m (– CA ) , 8 Apr 2016 (ex hort. Mar 2018 ), Phillips & Potgieter 34 ( NU0088343 !) . 2929 ( Underberg ): Cathkin Park , 1219 m (– AB ) , Jan 1953 , Bates 21 ( NU0010077 !) ; Cathkin Park , Ndema Forest , 1310 m (– AB ) , 3 Mar 1932 , Galpin 11826 ( PRE0036841 -0!); Cathkin Park , Ndema Forest , 1310 m (– AB ) , 3 Mar 1932 , Galpin s.n. ( BOL 50008!) ; Mpendle , Loteni River Valley , 1798 m (– AD ) , 13 Jan 1982 , Hilliard & Burtt 15083 ( NU0010092 !) ; Giants Castle Game Reserve , Forest Walk , (– BC ) , 4 Feb 1985 , Reid 974 ( NH0085048 -0!, PRE0676789 -0!); Kamberg Game Pass Farm (– BC ) , 11 Jan 1990 , Williams 711 ( NH0113530 -0!); Gxalingenwa Valley , Sani Pass road, 1630 m (– CB ) , 19 Feb 2015 , Braby 46 ( NU0044015 !) ; Gxalingenwa Valley , Sani Pass road, 1627 m (– CB ) , 19 Feb 2015 , Braby 76 ( NU0044054 !) ; Polela , Bamboo Mountain , 2134 m (– CB ) , Apr 1909 , Doidge 5572 ( K001208129 !) ; Bulwer , Enkelabantwana Forest (– CB ) , Mar 2000 , Edwards 2053 ( NU0010103 !) ; Cobham Forest Reserve , Emerald Vale , 1951 m (– CB ) , 4 Mar 1985 , Hilliard & Burtt 18303 ( E , K001208127 !, NU0010094 -0!); Cobham Forest Reserve , Sipongweni south side, 1890 m (– CB ) , 5 Mar 1983 , Hilliard & Burtt 16761 ( E , K001208128 !, NU0010093 !) ; Garden Castle (– CC ) , date unknown, Duckworth 168 ( CPF0021099 !) ; Mpendle , Everglades , 1524 m (– DB ) , 15 Oct 1964 , Moll 1265 ( NU0010084 !) ; Creighton , Lynn Avis Farm (– DD ) , Nov 1966 , Crewe 73 ( NU0010082 !) ; Creighton , Donnybrook , 1300 m (– DD ) , 4 Jun 2016 (ex hort. Mar 2018 ), Phillips 36 ( NU0088346 !) . 2930 ( Pietermaritzburg ): Mount West , 1672 m (– AA ) , 15 Apr 2017 , Phillips 57 ( NU0088392 !) ; Howick Falls (– AC ) , 4 Feb 1943 , Allsopp 69 ( NU0010068 !) ; Howick , Shafton (– AC ) , Mar 1900 , Hutton 7 ( GRA0005000 !) ; Howick , Umgeni Valley Nature Reserve , 973 m (– AC ) , 9 Mar 2016 (ex hort. Mar 2018 ), Phillips 23 ( NU0088330 !) ; Twin Falls , ± 25 km NNE of Howick , 365 m (– AD ) , 12 Feb 1988 , Grove 236 ( NU0010100 !) ; Karkloof , Caravan Bush Camp (– AD ) , 31 Jan 2012 , Martos 771 ( NU0049957 !) ; Karkloof , Bushwillow Park , 1356 m (– AD ) , 25 Aug 2016 (ex hort. Feb 2018 ), Phillips & Kiepiel 37 ( NU0088347 !) ; Karkloof , 914 m (– AD ) , Apr 1972 , Ripley 72 ( NU0010072 !) ; Karkloof (– AD ) , 20 Feb 1988 , Wirminghaus 555 ( NU0010096 !) ; Greytown (– BA ) , 4 Feb 1942 , Van der Merwe 2498 ( PRE0036812 -0!); Pietermaritzburg , Town Bush , 1006 m (– CB ) , Mar 1947 , Demont 34 ( NU0010074 !) ; Pietermaritzburg (– CB ) , Mar 1941 , Killick 13 ( NU0010071 !) ; Pietermaritzburg , Ferncliffe Nature Reserve , 762 m (– CB ) , 10 Feb 1979 , Nicholas 549 ( NU0010091 !) ; Pietermaritzburg , Ferncliffe Nature Reserve , 762 m (– CB ) , 10 Feb 1979 , Nicholas 550 ( NU0010089 !) ; Richmond , top of Hela Hela Pass (– CC ) , 25 Feb 1956 , Bell 14 ( BOL 49898!) ; Richmond , Enon Forest , 7 miles from Richmond to Byrne and Bulwer (– CC ) , 25 Feb 1956 , Bell 15 ( BOL 49897!) ; Richmond , Byrne , 1524 m (– CC ) , 31 Mar 1932 , Galpin 11993 ( K001208130 !) ; Richmond , Tala Farm , 914 m (– CD ) , 7 Mar 1966 , Moll 3056 ( NU0010080 !) ; Richmond (– CD ) , 11 Apr 1905 , Wylie s.n. ( NU0010059 !) ; Ingomankulu , ± 5 km SW of Camperdown , 973 m (– CD ) , 11 Jan 2016 (ex hort. Apr 2018 ), Young & Phillips 2473 ( BNRH !, NU0088296 !, NU0088297 !) ; Inchanga , 914 m (– DA ) , Apr 1953 , Bates 20 ( NU0010078 !) ; Table Mountain (– DA ) , date unknown, Killick 391 ( NU0010076 !) ; Monteseel (– DA ) , 31 Mar 1957 , Lawson 488 ( NH0047072 -0!); Inchanga , 609 m (– DA ) , 31 Mar 1957 , Loubser 46b ( NU0088548 !) ; Monteseel (– DA ) , 15 Mar 1964 , Strey 5204 ( NH0052546 -0!); Drummond (– DA ) , 6 Mar 1966 , Strey 6483 ( PRE0036825 -0!); Durban , Mgwahumbe Valley , 500 m (– DC ) , 11 Apr 1992 , Scott- Shaw 4653 ( CPF0021096 !) ; Krantzkloof Nature Reserve , 471 m (– DD ) , 13 Feb 2016 (ex hort. Mar 2018 ), Phillips & Garnier 14 ( NU0088528 !) ; Pinetown , Winston Park , 563 m (– DD ) , 15 Feb 2016 (ex hort. Mar 2018 ), Phillips 18 ( NU0088325 !) . 3029 ( Kokstad ): Umzimkulu , Ntsikeni Plateau , above Ntsikeni Plantation , 1775 m (– BA ) , 23 Jan 2002 , Bytebier et al. 2230 ( BR0000009353022 photo!, GRA0005021 !, NBG ); Kokstad , Zuurberg , 1067 m (– BC ) , Feb 1884 , Tyson 1683 ( BOL 50011!) ; Harding , Ingeli Forest , 977 m (– DA ) , 25 Jan 2016 (ex hort. Feb 2018 ), Grieve et al. 2003 ( NU0088294 !) ; Harding , Ingeli Forest , 983 m (– DA ) , 25 Jan 2016 (ex hort. Feb 2018 ), Grieve et al. 2004 ( NU0088295 !) . 3030 ( Port Shepstone ): Ixopo , Highflats , 887 m (– AC ) , 14 Jan 2016 (ex hort. Apr 2018 ), Young & Phillips 2504 ( BNRH !, NU0088302 !) ; Adams Mission , Nongwaan Falls , 152 m (– BA ) , 3 Apr 1945 , Adams 209 ( NU0010069 !) ; Umbumbulu , bottom of Nongwaan Falls (– BA ) , Apr 1945 , Lindahl 89 ( NU0010062 !) ; Umzinto , Vernon Crookes Nature Reserve , Hidden Valley , 420 m (– BC ) , 21 Nov 1984 , Balkwill & Cadman 2236 ( NU0010095 !) ; Dumisa , Ifafa , 500 m (– BC ) , 26 Mar 1908 , Rudatis 336a ( NH0106443 -0!, P00339733 photo!); Dumisa , Ifafa , 500 m (– BC ) , 5 Apr 1911 , Rudatis 1368 ( L1526648 photo!, P00339732 photo!, K001208131!, WAG1140774 photo!); Oribi Gorge Nature Reserve , Baboon Spruit (– CB ) , Feb 1973 , Davidson 2644 ( MO !) ; Izotsha Falls , view site, 457 m (– CB ) , 23 Feb 1964 , Hilliard s.n. ( BOL 49909!) ; Oribi Gorge Nature Reserve , 279 m (– CB ) , 6 Dec 2016 (ex hort. Mar 2018 ), Phillips & Kiepiel 39 ( NU0088349 !) ; Oribi Gorge Nature Reserve , 224 m (– CB ) , 6 Dec 2016 (ex hort. Mar 2018 ), Phillips & Kiepiel 40 ( NU0088378 !) ; Oribi Gorge (– CB ) , date unknown (ex hort. Feb 1966 ), Schelpe s.n. ( BOL 49903!) ; Umtamvuna Nature Reserve , Outeniqua Trail , 392 m (– CC ) , 11 Feb 2006 , Abbott 8371 ( PRU ) ; Umtamvuna Nature Reserve , Klipspringer , 384 m (– CC ) , 7 Jan 2016 (ex hort. Mar 2018 ), Grieve et al. 1977 ( BNRH !, NU0088290 !) . 3130 ( Port Edward ): Umtamvuma Nature Reserve , Clearwater Trails , 165 m (– AA ) , 20 Apr 2017 , Phillips 58 ( NU !) . Without precise locality: Umkomaas River , 1865, Gerrard 1536 ( BM000034620 !, K001208132 !) ; Natal , 1880, Wood 858 ( BOL 50014!) ; Zululand , 1865, Gerrard 1539 ( BM000034619 !, BR0000021817847 photo!, P00339736 photo!); Natal Highlands , Apr 1895 , Schlechter s.n. ( K001208134 !) . Limpopo : 2230 ( Messina ): Venda , 1 km from Dzamba en route to Rambuda (– CD ) , 12 Apr 1980 , Van Wyk 4101 ( PRU !) . 2329 ( Pietersburg ): Louis Trichardt , Hanglip (– BB ) , 25 Jan 1931 , Bremekamp & Schweickerdt 459 ( PRU !) ; Louis Trichardt , Hanglip Forest Reserve , 1462 m (– BB ) , 15 Mar 2017 (ex hort. Mar 2018 ), Phillips 54 ( BNRH !, NU0088390 !) ; Louis Trichardt , Zoutpansberg Mountains , about 5 miles above Louis Trichardt (– BB ) , 8 Mar 1948 , Rodin 4009 ( MO !) ; Houtboschberg , in jungle (– DD ) , Aug 1880 , Nelson 377 ( K001208122 !) . 2330 ( Tzaneen ): Duiwelskloof , Fairy Glen (– CA ) , 11Mar1958 , Scheepers 155 ( PRU !) ; Magoebaskloof , forest below Magoebaskloof Hotel , 1342 m (– CC ) , 12 Mar 2017 , Phillips 52 ( NU !) ; Magoebaskloof , Debengeni Falls , 1164 m (– CC ) , 17 Mar 2017 , Phillips 55 ( NU !) ; Woodbush [Houtbosh, Transvaal] (– CC ) , 1880, Rehmann 5856 ( BOL 50017!, K001208121 !) ; Letaba , Mtataspruit , just above provincial road-bridge, 945 m (– CC ) , 11 Mar 1958 , Scheepers 155 ( K001208126 !) ; Woodbush , near forester’s office, west slope (– CC ) , 4 Feb 1981 , Van Jaarsveld 6100 ( NBG0129535 -0!); Letaba (– CC ) , 27 Mar 1974 , Van der Walt s.n. ( BOL 49902!) . 2430 ( Pilgrim’s Rest ): Wolkberg , Lekgalameetse Nature Reserve , 1117 m (– AA ) , 13 Mar 2017 (ex hort. Mar 2018 ), Phillips 53 ( NU0088389 !) . Mpumalanga : 2430 ( Pilgrim’s Rest ): Bourke’s Luck Mine , on summit of Three Sisters mountain (– DB ) , 25 Feb 1937 , Galpin 14280 ( K001208123 !) ; Bourke’s Luck Mine , on summit of Three Sisters mountain , 1676 m (– DB ) , 25 Feb 1937 , Galpin s.n. ( BOL 49913!) ; Pilgrim’s Rest , Morgenzon S.F. (– DC ) , 8 Mar 1989 , Matthews 431 ( PRU !, NMB , ASU ) ; Graskop , God’s Window (– DD ) , 11 Feb 1998 , Goldblatt 10877 ( MO04928213 !) ; Graskop , Fairylands , next to Graskop Municipal Caravan Park , 1600 m (– DD ) , 29 Feb 1992 , Kurzweil 1657 ( NBG0150829 -0!); Graskop , God’s Window , 1680 m (– DD ) , 1 Feb 2016 (ex hort. Feb 2018 ), Phillips & Bytebier 6 ( NU0088315 !) ; Graskop , God’s Window , 1704 m (– DD ) , 1 Feb 2016 (ex hort. Feb 2018 ), Phillips & Bytebier 7 ( NU0088316 !) ; Graskop , God’s Window , 1659 m (– DD ) , 1 Feb 2016 (ex hort. Feb 2018 ), Phillips & Bytebier 8 ( BNRH !, NU0088317 !) ; Graskop , The Bonnet , 1636 m (– DD ) , 1 Feb 2016 (ex hort. Mar 2018 ), Phillips & Bytebier 9 ( NU0088318 !) ; Graskop (– DD ) , 11 Feb 1964 , Vorster & Coetzee 6 ( PRU !) . 2530 ( Lydenburg ): Dullstroom , Go Country Estate , 1755 m (– AC ) , 31 Jan 2016 (ex hort. Mar 2018 ), Phillips et al. 5 ( BNRH !, NU0088314 !) ; Sabie , Mac Mac Falls , 1249 m (– BB ) , 1 Feb 2016 (ex hort. Mar 2018 ), Phillips & Bytebier 10 ( BNRH !, NU0088319 !) ; Lydenberg , Spitskop (– BB ) , Apr 1887 , Wilms 1384 ( BM000034618 !, K001208124 !) ; Buffelskloof Nature Reserve , 1750 m (– BC ) , 18Feb 2012 , Martos 796 ( NU0049955 !) ; Buffelskloof Nature Reserve , 1750 m (– BC ) , 4 Feb 2016 (ex hort. Mar 2018 ), Phillips et al. 13 ( NU0088322 !) ; Lydenberg , Elandshoogte , 1900 m (– DA ) , 2 Mar 1978 , Richardson 106 ( NU0010090 !) ; Kaapschehoop Hiking Trail , 1675 m (– DB ) , 2 Feb 2016 (ex hort. Mar 2018 ), Phillips & Bytebier 11 ( NU0088320 !) ; Kaapschehoop , roadside between Kaapschehoop and Nelspruit , 1229 m (– DB ) , 2 Feb 2016 (ex hort. Mar 2018 ), Phillips & Bytebier 12 ( NU0088321 !) ; Nelsberg [ E . Transvaal] (– DD ) , 26 Feb 1936 , Taylor 1856 ( NY04157312 photo!); Nelsberg [ E . Transvaal ] (– DD ) , 26 Feb 1936 , Taylor 1857 ( BOL 49925!) . 2531 ( Komatipoort ): Barberton Mountains , near Havelock Mine , 1402 m (– CC ) , 13 Mar 1965 , Bayliss 2741 ( NBG0079970 -0!); Barberton (– CC ) , 22 Feb 1890 , Culver 23 ( BOL 50009!) ; Barberton Mountains , near Havelock Mine , 1500 m (– CC ) , 19 Feb 1979 , Linder 2050 ( BOL 49923!) . TANZANIA . T7. Iringa Region : Mufindi District , Nyalawa R. , 1981 m , 13 Apr 1971 , Paget-Wilkes 874 ( MO !) ; Udzungwa Mountains National Park , Mount Luhombero , 2040 m , 30 Sep 2000 , Luke et al. 6815 ( EA , K001208156 !) . Mbeya Region : Mbeya District , Poroto Ridge Forest Reserve , west of Lake Ngozi , 2103 m , 7 Mar 1970 , Wingfield 942 ( K000513090 !) ; Rungwe District , Rungwe Forest Reserve , south side, 1800 m , 28 Feb 1986 , Bidgood et al. 98 ( MO3440565 !) ; Rungwe District , Mwakaleli , vicinity of Mwatesi River , 9 May 1975 , Hepper & Field 5461 ( K000513093 !) ; Rungwe District , lava flow west of Kiejo Crater , 1800 m , 26 Mar 1958 , Pettersson et al. 388 ( EA , LIS , K000513091 !, NHT , UPS ) ; Rungwe District , Nyassa Highlands [Hochland], Station Kyimbila , 1912, Stolz 1973 ( L1526717 photo!, L1526716 photo!, U1475174 photo!, WAG1140772 photo!, WAG1140773 photo!) . Njombe Region : Lupembe area , upper Ruhudji River basin, north of the river, May 1931 , Schlieben 814 ( BR0000021817816 photo!, K001208155 !, P00339723 photo!) . ZIMBABWE . Manicaland Province : 1832 ( Umtali ) : Nyanga District , Placefell , 1829 m (– BC ) , 26 Mar 1949 , Chase 4124 ( BM000034607 !) ; Nyanga District , Romneydale , 1829 m (– BC ) , 22 Mar 1949 , Chase 4125 ( BM000034608 !) ; Nyanga District , forest ca. 5 km SE of Pungwe View , 1700 m (– BD ) , 9 Mar 1981 , Philcox et al. 8940 ( K001208150 !) . Without precise locality: Mutare [Umtali], 1585 m , 18 Feb 1926 , Eyles 4468 ( BOL 49916!) . 1932 ( Melsetter ): Vumba Mountains , Cloudlands , mountain slope east of Cripps Grid (– BA ) , 27 Feb 1949 , Chase 1375 ( BM000034605 !) ; Vumba Mountains , Castle Beacon , 1910 m (– BA ) , 18 Mar 1956 , Chase 6018 ( BM000034609 !) ; Vumba Mountains , Castle Beacon , 1830 m (– BA ) , 12 Mar 1981 , Philcox 8976 ( K001208149 !) ; Vumba Mountains Road , 10 miles from Mutare [ Umtali ] (– BB ) , 12 Feb 1968 , Ashton 151 ( GRA0005013 !) ; Vumba , Norseland , 1524 m (– BB ) , Mar 1949 , Wild 2802 ( BR0000021817823 photo!); Summit of Mount Peni , 1700 m (– DD ) , 16 Mar 1981 , Philcox et al. 8988 ( K001208148 !) . Without precise locality: Vumba Mountains , Mar 1931 , Woodburn NBG30/29 ( BOL 49884!) . 1933 ( Vila Pery ): Chimanimani Mountains , Stonehenge , 1829 m (– CC ) , 10 Feb 1958 , Hall 418 ( BOL 49892!) ; Chimanimani Mountains , south of Mount Peza , 1646 m (– CC ) , 2 Feb 1958 , Hall 297 ( BOL 96148!, K001208153 !) ; Chimanimani Mountains , forest behind mountain hut, 1737 m (– CC ) , 6 Apr 1969 , Simon & Kelly 1852 ( BOL 49924!, K001208152 !) . Masvingo Province : 2031 ( Bikita ) : Bikita District , south face of Mount Horzi , 1180 m (– BA ) , 11 May 1969 , Biegel 3136 ( BOL 49920!, K001208151 !) .