A taxonomic revision of the genus Ancylobothrys (Apocynaceae, Plumerioidae) in South Africa, including the description of a new geoxylic species

South African Journal of Botany 130 (2020) 117
122

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A taxonomic revision of the genus Ancylobothrys (Apocynaceae, Plumerioidae) in South Africa, including the description of a new geoxylic species R.A. Reddy*, K. Balkwill, F. Ralepele C.E. Moss Herbarium, School of Animal, Plant and Environmental Sciences, University of the Witwatersrand, Private Bag 3, WITS 2050, Johannesburg, South Africa

A R T I C L E

I N F O

Article History: Received 2 September 2019 Revised 6 December 2019 Accepted 14 December 2019 Available online xxx Edited by JS Boatwright Keywords: Ancylobothrys capensis Ancylobothrys petersiana Ancylobothrys pumilus Ancylobotrys Geoxylic suffrutex Maputaland endemic

A B S T R A C T

In South Africa, Ancylobothrys Pierre was last revised in 1963, although the taxa were maintained within the genus Landolphia P. Beauv. Later authors separated Ancylobothrys from Landolphia citing the tendril-like inflorescence and the softly pubescent fruit of Ancylobothrys as two of the most obvious characters separating these genera. A geoxylic form related to Ancylobothrys petersiana (Klotzsch) Pierre occurs in deep sandy soils of the Maputaland woody grassland in northern KwaZulu-Natal and is endemic to the Maputaland Centre of Endemism. This study shows that there is sufficient morphological distinction for this taxon to be considered a separate species, Ancylobothrys pumilus K. Balkwill & R.A. Reddy. The separation of A. pumilus from A. petersiana has necessitated a revision of the genus in South Africa. A key and descriptions to the South African species of Ancylobothrys are provided. © 2019 SAAB. Published by Elsevier B.V. All rights reserved.

1. Introduction Pierre (1898) created the genus Ancylobothrys from Landolphia for those taxa in Section Eulandolphia Pierre that were anatomically (open meristele, fruit lacking a sclerenchymous layer) and morphologically (large flowers, stamens located below the middle of the tube, a short style and an ovoid stigma) different. In 1909, Stapf placed Ancylobothrys back into Landolphia, but in Section Ancylobotrys, as he considered the differences between the taxa to be based on degrees of differences rather than discrete differences. According to Codd (1963), Pichon (1953) reinstated Ancylobothrys using the same arguments given by Stapf (1909). Codd (1963), however, maintained Stapf’s treatment as he considered it to be more reasonable. This circumscription was upheld in South Africa until 1992 when Persoon et al. published a revision of the genus Landolphia. According to Vonk et al. (1994), the tendril-like inflorescence and the softly pubescent fruit are two of the most obvious characters that separate Ancylobothrys from Landolphia. There has been considerable disagreement about the correct spelling for this generic name. The name was derived from the Greek words ancyla (hook) and botrys (grape) (Vonk et al., 1994). Pierre

* Corresponding author. E-mail address: [email protected] (R.A. Reddy). https://doi.org/10.1016/j.sajb.2019.12.021 0254-6299/© 2019 SAAB. Published by Elsevier B.V. All rights reserved.

(1898) originally, and incorrectly, wrote it as Ancylobothrys and in 1963 Huber corrected it to Ancylobotrys. This latter spelling has been commonly used. Article 60.1 of the International Code of Nomenclature for Algae, Fungi and Plants (Turland et al., 2018) states that a spelling should only be corrected if it was a typographical or orthographic error or if it needed to be standardized because of: letters and ligatures foreign to Latin; interchange between u and v, i and j or eu and ev; diacritical signs and ligatures; terminations; international Latinization; compounding forms; hyphens; apostrophes and full stops; or abbreviations. As Huber's (1963) name change does not meet any of these criteria, we follow Govaerts (1995 in WSCP). Codd (1963) observed that A. petersiana was very variable and that an erect, sparingly branched dwarf shrub form, with small, unbranched inflorescences occurs in the deep, sandy soils of northern KwaZulu-Natal (Zululand). We believe that there is sufficient evidence for this dwarf form to be elevated to species level. Ancylobothrys petersiana occurs in the Maputaland region of northern KwaZulu-Natal and eastern Limpopo (Codd, 1963). The typical form of A. petersiana is a climber that occurs in forests or along forest margins. The new taxon with its erect, sparingly branched stems and often unbranched inflorescences (Codd, 1963), is restricted to the endemic Maputaland Woody Grassland (CB2) of the Indian Ocean Coastal Belt Biome (Mucina and Rutherford 2006; Siebert et al., 2011) in northern KwaZulu-Natal which also constitutes the

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Maputaland Centre of Endemism. This unique grassland is sandy and has a shrubby appearance; due to the many geoxylic shrubs that occur there (van Wyk and Smith 2001; Mucina and Rutherford 2006). Ancylobothrys capensis (Oliv.) Pichon is uncommon in KwaZuluNatal; only one specimen, Mogg 7615, from Newcastle (2729 DB), is recorded on the SANBI new POSA website and is also the only specimen cited by Codd (1963) as occurring in this province. This species occurs in dry, bush-covered rocky areas, looks superficially similar to A. pumilus, but differs from the other two species in having smaller leaves, secondary veins closer together and the angle between the secondary and mid veins are close to 90° Currently the genus consists of seven species worldwide with two occurring in South Africa (BODATSA, 2019). Both the separation of A. pumilus from A. petersiana, and because in the last revision Codd (1963) still placed the species of Ancylobothrys in the genus Landolphia, a revision of the genus in South Africa is needed. This account recognizes three species in South Africa, and provides a revised key and circumscriptions for all three taxa that occur in South Africa. 2. Material and methods Herbarium specimens at J, NH, NU and PRE were consulted (acronyms after Thiers, 2019). Digital images of specimens from these herbaria, as well as from K (on JSTOR Global Plants), were measured using the Digimizer Image Analysis Software program Version 4.6.1 (https://www.digimizer.com). Field observations were also made. Species distributions were plotted in DIVA-GIS version 7.5 (http:// www.diva-gis.org/gdata) using label information from specimens as well as from localities listed in the Botanical Database of Southern Africa (BODATSA, 2019). The additional specimens examined for each species are recorded using the Quarter Degree Reference System of Edwards and Leistner (1971).

very prominent; pulp edible. Seeds few or many, embedded in pulp, endosperm cartilaginous. A genus of 7 species in Africa; with one species extending to Madagascar and the Comores. In South Africa, the genus is confined to the north-eastern part of the country (Fig. 1). 3.2. Key to the South African species

1. Woody climber; inflorescences
60 mm long; corolla lobes narrower than 3 mm; styles 1 mm long . . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .2. A. petersiana Scandent or geoxylic shrub; inflorescences  60 mm long; corolla lobes > 3 mm wide; styles 1.5 mm long . . .. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . 2 2. Geoxylic shrub with erect aerial stems, rarely > 0.5 m high; leaves longer than 35 mm, angle between main and secondary veins on leaves acute, secondary veins 2.4
4.8 mm apart . . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .3. A. pumilus Low growing or scandent shrub, to 3 m high; leaves to 35 mm long, secondary veins on leaves appear § rectangular, secondary veins 1.5
3.0 mm apart. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .1. A. capensis

3.3. Taxonomic descriptions
m. Inst. Fran¸c . Afrique 1. Ancylobothrys capensis (Oliv.) Pichon in Me Noire 35: 297 (1953). Landolphia capensis Oliv. in Hook. Ic. Pl. 13: t. 1228 (1877); Stapf in Fl. Cap., 4, 1: 495 (1907); Phillips in Fl. Pl. South Africa 9: t. 352 (1929); Marloth in Fl. S. Afr. 3: t. 17 (1932);

3. Taxonomy 3.1. Genus description
r. 2,1: 91 Ancylobothrys Pierre in Bull. Mens. Soc. Linn. Paris se (1898); Hua in Bull. Mus. Hist. Nat. (Paris)5: 186 (1899); Pichon
m. Inst. Fran¸c . Afrique Noire 35: 272 (1953). Ancylobotrys Huber in Me in Fl. W. Trop. Afr. 2nd ed. 2: 60 (1963); Vonk et al. in Wageningen Agric. Univ. Pap. 94 (3): 1
44 (1994). Landolphia P.Beauv in Fl. Oware 1: 54, t. 34 (1806) pro parte; Stapf in Fl. Cap. 4(1): 492
496 (1907) pro parte. Type species: Ancylobothrys petersiana (Klotzsch) Pierre, designated here. Perennial shrubs, woody climbers, or dwarf shrubs; milky latex present. Leaves dark green above, paler below, simple, entire, opposite, leathery, variable in shape from elliptic through oblong to obovate, midrib channeled above, raised below, pinnately nerved; petioles short; stipules absent. Inflorescences of long terminal panicles, often acting as tendrils. Flowers hermaphroditic, pentamerous, sweetly scented, small to medium-sized, sessile or pedicelled. Calyx united at the base; lobes triangular, imbricate, bright green, small, leathery, eglandular, usually densely rusty pubescent. Corolla tube pink with white base, narrowly cylindrical, swollen just above the base, with long hairs within tube, usually pubescent on outer surface; lobes 5, overlapping to the left, usually longer than the tube, occasionally fringed with white hairs, white, often tinged pinkish; stamens inserted in the lower half of the corolla tube. Androecium anthers 5, bi-thecous, introrse, with inward facing longitudinal slits, ovate, apices acute; filaments very short, epipetalous. Gynoecium stigma level with the base of anthers, bifid, arms converging; style glabrous, terete, short, stout; ovary superior, unilocular, hairy; ovules many, placentation parietal. Fruit a many-seeded, globose berry, with a hard rind; wall not sclerified; surface densely velvety; placenta

Fig. 1. Distribution of Ancylobothrys species in South Africa.

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Verdoorn in Edible Wild Fr. Transv. Bull. 185: 45, t. 7 (1938); Miller in Journ. S. Afr. Bot. 18:71 (1952). Type: South Africa, Transvaal, W. of Pretoria, McLea 3098 (K000233580
Global Plants!, lecto., designated by Pichon, 1953; BM000925633
Global Plants!, isolecto.). Much branched low growing or scandent shrub, to 3 m high, often forming a dense bush. Stems with many small, often raised lenticels, fine rusty hairs when young, older stems glabrescent. Leaves 22
35 £ 10
18 mm, midrib shallowly channeled above, secondary veins, 1.5
3.0 mm apart, spreading at almost right angles to the midrib, pubescent on both surfaces when young, becoming glabrescent with age; apex rounded; base obtuse; petiole 3
7 mm long. Inflorescences in terminal panicles, 24
100 mm long, ending in 1
3-flowered cymes, on short, spreading or recurved branches which often act as tendrils, rusty pubescent to glabrescent; bracts small, triangular, with white or rusty hairs, sometimes with white hairs at tips. Calyx 2.5
5.0 £ 2.0
3.0 mm; tube 1
3 mm long; lobes 2
4 mm long. Corolla tube 5
10 mm long, swollen to 1
2 mm wide slightly above base, then narrowing again before widening to 1
2 mm at throat; lobes white, 6
16 £ 3
5 mm. Androecium anthers c. 1.5
2.5 mm long; filaments c. 1
1.5 mm long. Gynoecium stigma at base of anthers, apex bifid; style cylindrical, c. 1.5 mm long; ovary globose, pubescent; ovules small, numerous. Fruit globose to pearshaped; densely and finely reddish tomentose when young, yellowish when ripe; 35
50 mm in diameter. Seeds few to many. Distribution and ecology: A. capensis is widely distributed and has a more inland distribution in southern Africa; occuring in Mpumalanga, Gauteng, Limpopo, North West Province and eastern Botswana. There is only a single collection known from north-western KwaZulu-Natal. It is frequently found in rocky areas in grassland and bushveld. Phenology: Flowering August
December peaking in October. Fruiting is mainly in January. Additional specimens examined: South Africa. LIMPOPO. 2428 (Nylstroom): on the farm Nooitgedacht, 1200 m (
CD), 3 Dec 1986, Balkwill & Cadman 3664 (J). NORTH WEST. 2526 (Zeerust): Marico Dist. (
CB), Dec 1926, van der Merwe 27 (PRE); Rustenburg Dist; Zwartruggens; deep gorge in tributary of Elandsriver (
DB), 6 Dec 1934, Sutton 894 (PRE). 2527 (Rustenburg): on the farm Boshoek 103 JQ, 1600 m (
AC), 24 Aug 1943, Rose Innes 22 (J, PRE); Grootkloof, near Hartebeespoort, 1800 m (
DB), 10 Oct 1988, Murray 14 (J); Rustenburg Dist. Magaliesberg (
CA), 3 Oct 1905, Pegler 965 (PRE); Rustenburg Dist.; Boekenhoutfontein Farm (
CA), 14 Sep 1943, RoseInnes 78 (PRE); Rustenburg (
CA), Jan 1935, MacGregor 384 (PRE); Magaliesberg, Between Maanhaarand and Buffelspoort Dam, on the Farm Kloofwaters, 1400 m (
CD), Laburn 95 (J); Brits Dist., Jacksonstuin (
DA), 19 Sep 1957, van Vuuren 277 (PRE); Hartebeespoort Nek (
DD), 10 Oct 1949, Prosser 1298 (PRE). MPUMALANGA. 2529 (Witbank): Middelburg Dist., Olifants River Gorge, “20 radial mi.”[32 km] N.W. of Johannesburg, on the farm Slagthoek 250 (126) JS, 1200 m (
CA), 10 Oct 1953, Mogg et al. 22,478 (J, PRE). 2630 (Carolina): Carolina (
AA), Nov 1917, Leipoldt TRV 18,639 (PRE). GAUTENG. 2527 (Rustenburg): Hekpoort, Timeball Hill, Uitkomst 499 JQ Farm (
DC), 29 Mar 1971, Coetzee 685 (PRE); Magaliesberg, De Kloof (
DD), 21 Jan 1967, Dahlstrand 1915 (PRE). 2528 (Pretoria): Magaliesberg, Pretoria Dist., Wonderboom Res. (
CA), 24 Aug 1944, Repton 1795 (PRE); Pretoria Dist., Near Pretoria, Baviaanspoort (
CB), 25 Sep 1925, McClean 61 (PRE); Daspoort Range (
CC), Oct 1924, Kaplan TRV 26,071 (PRE). 2627 (Potchefstroom): on the farm Waterval (74)174 IQ, 1750 m (
BA), 25 Oct 1950, Mogg et al. 20,284 (J); Northcliff. Ridge at the corner of Rockey and Louie Streets, 1550 m (
BB), 12 Oct 1992, Hartley, Reddy & Masilo 2035 (J); Witpoortjie Kloof & Waterfalls (Walter Sisulu Gardens), gorge, 1650 m (
BB), 4 Oct 1917, Moss 4693 (J); on the farm Olifantsvlei (16) 327 IQ, 1750 m (
BD), 14 Oct 1950, Mogg et al. 20,213 (J); Losberg, Potchefstroom Dist., Elandsfontein (
DA), 20 Dec 1934, Theron 1001 (PRE). 2628 (Johannesburg): Melville Koppies Nature Reserve, 1800 m (
AA), 6 Feb 1991,

119

House 16 (J); 3.2 km W of Alberton, Klipriviersberg 106 IR, 1700 m (
AA), 7 Jan 1950, Mogg 18,260 (J); Lyme Park, off Nichol highway, 1550 m (
AA), 2 Dec 1973, Munday 638 (J); Mondeor Hills, 1650 m (
AC), Patterson s.n. sub J 37,810 (J); Heidelberg Kloof, 1650 m (
AD), 3 Jan 1953, Mogg et al. 20,450 (J); Springs Dist., Waaikraal (
AD), Feb 1938, Naude 6562 (PRE); Heidelberg (
AD), Nov 1909, Leendertz TRV 7702 (PRE); Suikerbosrand/Sugarbush Mountains, 1700 m (
CA), 15 Oct 1949, Mogg 18,089 (J); Suikerbosrand, Perdekop (
CA), 25 Sep 1971, Bredenkamp 36 (PRE); 8 km SW of Heidelberg, Sedaven Koppie, 1600 m (
CB), 13 Nov 1985, Hartley 421 (J); Rooikoppies, S. of Heidelberg on rd. to Villiers (
CB), 23 Oct 1938, Gillett 4610 (PRE). KWAZULU-NATAL. 2729 (Volksrust): Allerdene (
DB), 10 May 1920, Mogg 7615 (PRE). 2. Ancylobothrys petersiana (Klotzsch) Pierre in Bull. Mens. Soc. Linn.
r. 2,1: 91 (1898); Pichon in Me
m. Inst. Fran¸c . Afrique Noire Paris se 35: 290 (1953). Willughbeia petersiana Klotzsch in Naturw. Reise Mossambique [Peters] 6 (Bot., 1): 281 (1861). Landolphia petersiana (Klotzsch) Dyer ex Hook. f. in Kew Gard. Rep. 1880: 42 (1881); Dyer in Hooker's Icon. Pl.28: t. 2756(1903); Stapf in Fl. Trop. Afr. [Oliver et al.] 4, 1: 47 (1902); Stapf in Fl. Cap.4, 1: 493 (1907); Codd in Fl. southern Afr. 26: 260 (1963). Pacouria petersiana (Klotzsch) S. Moore in Journ. Linn. Soc., Bot. 37: 180 (1905). Types: Mozambique, Sofala, Sena, Peters s.n. (By, holo.); Niassa, Antonio Enes (= Angoche), Gomes e Sousa 4856 (K000233589
Global Plants!, neo., designated by Vonk et al., 1994; COI, isoneo.; PRE, isoneo.; WAG0000102
Global Plants!, isoneo.). Willughbeia senensis Klotzsch in Peters, Naturw. Reise Mossambique [Peters] 6 (Bot., 1): 281 (1861). Landolphia senensis (Klotzsch) K. Schum. in Pflanzenw. Ost-Afr. B: 453 (1895). Type: Mozambique, Sofala, Sena, Peters s.n. (By, holo.). Landolphia monteiroi N.E. Br. in Monteiro, Delagoa Bay, 161, 163, 178 (1891); Dyer ex Stapf in Bull. Misc. Inform. Kew 1907(2): 51 (1907); Codd in Mem. Bot. Surv. S. Afr. 26: 155 (1951). Type: Delagoa Bay, Monteiro 37 (K000233584
Global Plants!, holo.). Landolphia angustifolia K.Schum. ex Engl. in Abh. Preuss. Akad. Wiss. Berl. Phys.-Math. Kl., 1:34 (1894). Landolphia scandens var. angustifolia (Engl.) Hallier f. in Jahrb. Hamb. Wiss. Anst. 17, Beih. 3: 84 (1900). Landolphia petersiana var. angustifolia (Engl.) Stapf in Fl. Trop. Afr. 4, 1: 48 (July 1902). Type: Tanzania, Usambara Mountains, Holst 2220 (K000233594
Global Plants!, lecto., designated by Vonk et al., 1994; HBG-502,719
Global Plants!, isolecto.; M-0,110,097
Global Plants!, isolecto.).
vre in Ann. Soc. Sci. Landolphia petersiana var. rotundifolia Dewe vre) Pierre Bruxelles 19: 122 (1895). Ancylobotrys rotundifolia (Dewe
r. 2, 1: 92 (1898). Landolphia scandens in Bull. Mens. Soc. Linn. Paris se vre) Hallier f. in Jahrb. Hamb. (Schum.) Didr. var. rotundifolia (Dewe Wiss. Anst. 17, Beih. 3: 82 (1900). Type: Tanzania, Zanzibar, Joblonsky 23 (P, holo.) Landolphia scandens var. stuhlmannii Hallier f. in Jahrb. Hamb.  sch, Wiss. Anst. 17, Beih. 3: 83 (1900). Type: Tanzania, Amboni, Bu Holst 2563 (HBG 502,720
Global Plants!, lecto., designated by Vonk et al., 1994; M-0,110,096
Global Plants!, isolecto.). Landolphia scandens var. tubeufii Busse in Bot. Jahrb. Syst.32: 171 (May 1902). Type: Kenya, Busse 1051 (G00015120
Global Plants!, lecto., designated by Vonk et al., 1994; B100154162
Global Plants!, iso.; HBG-502,721
Global Plants!, iso.). Landolphia petersiana var. rufa Stapf in Fl. Trop. Afr. 4, 1: 48 (July 1902). Syntypes: Malawi, “Nyasaland’’, 1891, Buchanan 437 (BM; K000233585
Global Plants!, lecto., designated by Vonk et al., 1994); Mozambique, Tete, xii.1858, Kirk s.n. (K000233588
Global Plants!, syn.); Tete, ii.1859, Kirk s.n. (K000233586
Global Plants!, syn.). Woody climber. Stems many, seldom branched, young stems with fine rusty or white hairs, older stems dark brown with white hairs, becoming glabrescent, hairs long, crisped, with many small, often

120

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raised lenticels. Leaves 49
80 £ 16
30 mm, with white hairs above when very young, especially at apex, becoming glabrescent with age, rusty pubescent below when very young, especially on midrib and at apex, also becoming glabrescent with age; base cuneate to slightly rounded; apex variable (rounded, acuminate or acute); veins light to yellowish green above, whitish to cream below, secondary veins dark green below, 3
6 mm apart, angle between the secondary and mid veins 60°; margins slightly revolute, yellowish; petiole 4
8 mm long, grooved above with white hairs. Inflorescence paniculate, 60
115 mm long, flowers sessile, clustered at ends of branches, often acting as tendrils; bracts small, triangular, with rounded apices, white hairy, sometimes with rusty hairs basally. Calyx 3
8 £ 2
3 mm; tube 1.0
2.5 mm long; lobes 1
3 mm long. Corolla tube 7
12 mm long, swollen to 1.5
2.0 mm wide slightly above base, then narrowing again before widening to 2
3 mm at throat; lobes white without, sometimes pinkish within with a dark pink midline, margins with long white hairs, sometimes only sparsely so, 12.0
20.0 £ 2.5
3.0 mm. Androecium anthers c. 2 mm long; filaments c. 2 mm long. Gynoecium stigma c. 2 mm long, apex bi-lobed; style c. 1 mm long; ovary densely white pubescent; ovules small, numerous. Fruit globose, velvety pubescent, 30
60 mm in diameter. Seeds whitish, flattened. Nomenclatural notes: Both Leeuwenberg and Kupicha (1985) and Vonk et al. (1994) credit Stapf (1902) with the combination Landolphia petersiana var. tubeufii Busse ex Stapf. However, Stapf only mentions the name Landolphia scandens var. tubeufii (as var. tubeufii) in his notes at the end of Landolphia petersiana and does not make any € cher in combination. In addition, the fascicle of Botanische Jahrbu which Busse published appeared in May 1902 whereas the fascicle of Flora of tropical Africa in which Landolphia appeared was published in July 1902, thus the authorship of the names as indicated by Leeuwenberg and Kupicha (1985) and Vonk et al. (1994) is incorrect and the combination, Landolphia petersiana var. tubeufii, has never been created. The isotypes of Landolphia scandens var. tubeufii located at B and HBG are fragmentary and so although Busse worked at B, we agree with Vonk et al. (1994) in designating the sheet at G as the lectotype. The precise dates of publication of Volume 32, Fasicle 1 of Bot€ cher and Volume 4, Section 1, Fascicle 1 of Flora of anische Jahrbu tropical Africa mean that Landolphia scandens var. tubeufii has priority over Landolphia petersiana var. rufa. Distribution and ecology: A. petersiana occurs on the eastern part of the African continent from Somalia to South Africa as well as in Madagascar and the Comores. In Madagascar it is restricted to the north-eastern coastal dunes (Vonk et al., 1994). Phenology: Flowering August
February, fruiting sporadic, beginning in November and peaking in February. Additional specimens examined: South Africa. LIMPOPO. 2229 (Waterpoort): About 40 km WNW of Louis Trichardt and 28 km ENE of Vivo, Western Soutpansberg, Leshiba Wilderness Area, Hamasha Bush Camp, 1164 m (
DC), 16 Nov 2016, Bester 13,518 (PRE); Zoutpansberg, Wyllie’s Poort, (
DD), 16 Oct 1956, Story 5944 (PRE). 2230 (Messina): Musina (
BD), 29 Feb 1984, Botha 3416 (PRE); Venda, Masisi, 379 m (
BD), 23 Sep 1979, Netshiungani 942 (J, PRE). 2231 (Pafuri): Behind Punda Maria, 500 m (
CA), 31 Oct 1948, Codd & Dyer 4548 (PRE); Dzundweni Hill, 3 km SE of Punda Maria, 500 m (
CA), Nov 1948, Codd & Dyer 4592 (PRE). KWAZULU-NATAL. 2632 (Bela Vista): Ndumo Game Reserve, c.85 m (
CD), 8 Nov 1989, Ward 2683 (PRE); Ndumo Hill (
CD), Pooley 137 (NH); 10 km W of Muzi Police Station, 60 m (
DC), 23 Oct 1971, Moll & Nel 5580 (NH); Makanes Pont, Mengwu sand forest (
DC), 13 Nov 1968, Venter 5169 (PRE); Maputoland, Kosi Bay, west of the reserve, along the road (going) in (
DD), 16 Sep 1990, McMurtry & Burns 8586 (PRE); Kosi Bay (
DD), 9 Sep 1971, Strey 10,430 (NH); Near Kosi Nature Reserve, Kosi Bay, <90 m (
DD), 24 Sep 1961, Edwards 2548 (NU, PRE); Tongaland, between tar road and Kosi mouth entrance gate, 36 m (
DD), 13 Oct 2010, Meyer 5185 (PRE). 2732 (Ubombo): 8 km W of Manzengwenya thicket/grassland, c. 30 m (
BA), 24 Feb 1971, Moll 5218 (NH,

PRE); Bhanga Nek (
BB), 6 Dec 1980, Gwala 206 (NU); Lalanek (
BC), 3 Oct 1984, Botha 3454 (PRE); Eziqeni area, c. 20 m (
BC), 27 Jan 1983, Ward 355 (NH); Lake Sibaya National Park, § 12 km NNW of Mabibi, E of lake, 31 m (
BC), 9 Feb 2018, Bester 14,109 (PRE); Mkuze Game Reserve, c. 80 m (
CC), 26 Aug 1975, Goodman 313 (NU); Mtante Sand Forest, 100 m (
CD), 5 Jul 1985, Balkwill, Cadman & Stormans 3056 (J); Sodwana Bay National Park, c.60 m (
DA), 9 Oct 1960, Ward 3497 (PRE); Sodwana Bay Nat. Park/IsimangalisoNat. Park, 15 m (
DA), 26 Aug 2016, Balkwill & Balkwill 13,841 (J); Sodwana State Forest, 40 m (
DC), 24 Sep 1987, MacDevette 1783 (NH). 2832 (Mtubatuba): Nyalazi State Forest (
AB), 1 Oct 1984, Gordon 199 (NH); Lake View (
AC), 23 Sep 1943, West 2098 (NH, PRE); Palm Ridge Farm, 60 m (
AC), 12 Sep 1967, Harrison 104 (NH, PRE); St. Lucia Estuary, 100 m (
AD), 21 Sep 1981, Hunter 298 (J); St. Lucia System, Lake Bhangazi, 15 m (
BA), 15 Feb 1997, Ward 13,916 (PRE). 3. Ancylobothrys pumilus K. Balkwill & R.A. Reddy sp. nov. Type: South Africa, KwaZulu-Natal, 2732 (
DA): Northern KwaZuluNatal, uMkhanyakude District, Sodwana Bay National Park/ IsimangalisoNational Park, [27°340 27.200 S 33°390 7.700 E], 25 m (
DA), 26 Aug 2016, K. Balkwill & L.C. Balkwill 13,827 (J, holo.; B, K, M, MO, PRE, RSA, iso.). Geoxylic shrub, underground stems branching, spreading; aboveground stems mainly erect, 0.3
0.5 m high, occasionally creeping, with lenticels often obsured by white or rusty hairs. Leaves 36
59 £ 14
26 mm, midrib whitish to cream above, secondary veins light to yellowish green above, dark green below, 2
5 mm apart, angle between the secondary and mid veins 60°; with white hairs on dorsal surface when very young, especially at apex, becoming glabrescent with age, rusty pubescent below when very young, especially on midrib and at apex, also becoming glabrescent with age; apex variable (rounded, acuminate or acute); margins slightly revolute, yellowish; base cuneate to slightly rounded; petiole 4
7 mm long, with white hairs, grooved above. Inflorescences in few-flowered axillary cymes, 16
50 mm long, only occasionally acting as tendrils, white pubesecent, sometimes with rusty hairs basally; bracts small, triangular, with rounded apices. Calyx 3.0
6.0 £ 2.0
3.5 mm; tube 1.5
3.0 mm long; lobes 2
3 mm long. Corolla tube 16
31 mm long, swollen to 1
3 mm wide slightly above base, then narrowing before widening to 2
6 mm at throat; lobes white without, white or sometimes pinkish with a dark pink midline within, 16
31 £ 3.5
6 mm; margins with long white hairs, sometimes only sparsely so. Androecium anthers c. 2 mm long; filaments c. 2 mm long. Gynoecium stigma c. 2 mm long, apex bi-lobed; style c. 1.5 mm long; ovary densely white pubescent; ovules small, numerous. Fruit globose, velvety pubescent, 16
25 mm in diameter. Seeds whitish, flattened. Etymology: Named for its habit — pumilus is Latin for “dwarf”. Diagnostic characters: Ancylobothrys pumilus differs from A. petersiana in having slightly shorter, narrower leaves (36
59 £ 14
26 mm, not 49
80 £ 16
30 mm as in A. petersiana), much shorter inflorescences (16
50 mm rather than 60
115 mm), which rarely act as tendrils; larger flowers (16
31 £ 3.5
6 mm compared to 12.0
20.0 £ 2.5
3.0) and a slightly longer style (1.5 mm versus 1.0 mm long) (Figs. 2 and 3). In addition, A. pumilus is an underground shrub with stems 0.3
0.5 m high that is endemic to the Maputaland Woody Grassland, whereas A. petersiana is a woody climber or scandent shrub with stems longer than 1 m, is more widespread and occurs in woodland, thickets and forest margins. A. capensis has smaller leaves (22
35 £ 10
18 mm) than A. pumilus (36
59 £ 14
26 mm), with secondary veins closer together and the angle between the secondary and mid-veins appearing to be close to 90° (rather than 60° as in A. pumilus). The distribution of A. capensis does not overlap with that of the new species (Fig. 1). Distribution and ecology: confined to, but widespread, in the Maputaland Woody Grassland (CB2) (Mucina and Rutherford 2006)

R.A. Reddy et al. / South African Journal of Botany 130 (2020) 117
122

Fig. 2. Ancylobothrys petersiana showing the climbing habit (A); flowers and tendrillike inflorescences (B). (Images: K. Balkwill and D. Oosthuizen). Scale bars = 10 mm.

of the Indian Ocean Coastal Belt Biome. The distribution of A. pumilus overlaps with that of A. petersiana; it is however confined to the deep sandy grassland rather than occuring in forest and at forest margins. Phenology: Flowering July
January, fruiting sporadic, beginning in October. Conservation notes: While A. pumilus appears to be restricted to the vulnerable Maputaland Centre of endemism, it is fairly well distributed with an area of occurrence of more than 60 000 km2 and

121

many of the localities occur in protected reserves. This is a long lived, deep rooted perennial species with well-established populations. Based on the wide extent of occurrence and the number of protected locations in which this species is found, we recommend a provisional threat status of least concern (LC) (IUCN, 2012). Additional specimens examined: South Africa. KWAZULU-NATAL. 2632 (Bela Vista): Near Maputa Store, 30.5 m (
DD), 25 Nov 1967, Strey & Moll 3888 (NH); Kosi Bay (
DD), 9 Sep 1971, Strey 10,427 (NH, PRE); 2 km from Natal Parks board Camp to Kosi Mouth (
DD), 9 Sep 1971, Stephen 363 (PRE). 2732 (Ubombo): NE of Sihangwane Store, about 99 m (
AB), 5 Jan 1964, Tinley 917 (NH); Western border fence of Tembe Elephant Park, 118 m (
AB), 14 Aug 2002, Meyer 4108 (PRE); Western border fence of Tembe Elephant Park, 97 m (
AB), 20 Aug 2002, Meyer 4127 (PRE); Makatini Flats (
AC), March 1967, Van der Merwe s.n. (PRE); Near Mabibi (
BA), 29 Oct 2001, Potgieter 529 (NU); Road to E tip of Lake Sibayi (
BA), 10 Dec 1968, Pooley 253 (NU); Maputaland, Sileza (
BA), 7 Nov 1992, Williams 972 (NH); KwaZibi River crossing, 305 m (
BA), 15 Sep 1970, Ross & Moll 5061 (NH); 8 km due S of Maputa, 30.5 m (
BA), 26 Nov 1969, Moll 4710 (NH); Approx 4 miles [6.4 km] north of Ngutshana Pans on Maputa Rd, c. 76 m (
BA), 11 Sep 1961, Ward 3725 (NH, PRE); Siyandla Forest, c. 3 km from control gate (
BB), 19 Jul 1986, Ngwenya 309 (NH); Black Rock (
BB), 3 Oct 1984, Botha 3466b (PRE); Othungweni area, ca 35 m (
BC), 26 Aug 1982, Ward 229 (NH); Maputaland, Manzengwenya (
BC), 8 Nov 1992, Williams 977 (NH); 1 km from Lake Sibaya, on the road to Mbaswane (
BC), 26 Nov 1975, Botha & van Wyk 1161 (PRE); Sodwana Bay NationalPark/IsimangalisoNational Park (
DA), 15 Jun 2016, Balkwill, McCallum & Oosthuizen 13,784 (J); 26 Aug 2016, Balkwill & Balkwill 13,821 (B, J, K, MO, PRE); Balkwill & Balkwill 13,828 (J); 14 Nov 2016, Balkwill, McCallum, Reddy, Oosthuizen & Balkwill 13,854 (J); Sodwana State Forest (
DA), 6 Oct 1986, 40 m, MacDevette 1239 (NH); Sodwana State Forest, c. 30 m (
DA), 6 Jul 1986, Nicholas, Bishop & Perks 1455 (NH); (M)bazwana (
DA), 22 Oct 1938, Gerstner 3694 (NH); 11 Jan 1964, Strey 5121

Fig. 3. A. pumilus showing a pink tinged flower (A), the large, velvety fruit (B) and habit (C). (Images: K. Balkwill (A), L.C. Balkwill (B) and J. Burrows (C)). Scale bars = 10 mm.

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122

(NH); Maputaland (
DA), 15 Aug 1969, Phillips s.n. sub NU 47,016 (NU). 2832 (Mtubatuba): 16 miles (25 km) N of Matubatuba (
AC), 14 Oct 1946, Acocks 13,084 (PRE); Codd 2015 (PRE); Eastern Shores, Lake St Lucia (
AD), Sep. 1973, Smook 486 (NU); St Lucia Game Park, 2 Sep. 1983 (
AD), McLean 24 (NH); E. of Lake St. Lucia, near “The Wathe”, c. 12 m, (
BA), 30 Oct 1968, Ward 6684 (PRE); Richards Bay, Alton (
CA), 9 Oct 1995, Hutchings & Williams 3552 (NH); Richards Bay (
CC), 16 Apr 1939, Gerstner 3758 (NH); Richards Bay (
CC), 1 Dec 1938, Gerstner 4600 (PRE). Other possible localities reported in publications are: Sokhulu Forest and Mapelane Nature Reserve, near St Lucia (2832AD) (Grainger, 2011) and west of Richard’s Bay (2831DD) (Siebert et al., 2011).

Declaration of Competing Interests There are no interests to declare.

Acknowledgements We thank CREW (Custodians of Rare and Endangered Wildflowers) for organizing the trips on which we first saw the new species in the field. We are grateful to Isimangaliso Wetland Park and Ezemvelo KZN Wildlife for granting us a collecting permit (permit number: OP 38/2016). We thank the curators of NH, NU and PRE for the use of their specimens and images. We are grateful to the National Research Foundation (Grant Number 119295) and the University of the Witwatersrand for financial support. We thank Donald McCallum for assisting with the graphics. We appreciate the assistance in the field and photographs provided by Lindsay Carolyn Balkwill, John Burrows and Delia Oosthuizen. Finally we gratefully thank the reviewers for their helpful comments and insights.

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