Taxonomy of the Southern African Psoralea Aphylla Complex (Psoraleeae, Leguminosae)

South African Journal of Botany 97 (2015) 77–100

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South African Journal of Botany

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Taxonomy of the southern African Psoralea aphylla complex (Psoraleeae, Leguminosae)

Meshack N. Dludlu ⁎, A. Muthama Muasya, Samson B.M. Chimphango, Charles H. Stirton

Bolus Herbarium, Department of Biological Sciences, University of Cape Town, Private Bag X3, Rondebosch 7701, South Africa article info abstract

Article history: The genus Psoralea L., comprising about 70 species, has its centre of diversity and endemism in the Greater Cape Received 22 April 2014 Floristic Region of South Africa. Species delimitation and identification in the genus are difficult. Many putative Received in revised form 11 November 2014 species are unpublished or known only by manuscript names. As a first step to address this problem this study Accepted 27 November 2014 reports on the taxonomy of the Psoralea aphylla complex — a distinct group of species characterised by either a Available online xxxx complete absence of leaves, a reduction of the leaves into scales, or the occurrence of small filiform leaflets. Plants Edited by GV Goodman-Cron may be suffrutices, small to large shrubs, or trees. Morphological and ecological analyses of herbarium specimens and recent field collections distinguished ten species within the complex. Three species (P. aphylla L., P. filifolia Keywords: Eckl. & Zeyh. and P. peratica C.H.Stirt.) are already published, while the seven other taxa have been known by Leguminosae manuscript names and are formalised in this paper (P. congesta C.H.Stirt. & Muasya; P. fleta C.H.Stirt.; P. gigantea Psoralea aphylla complex Dludlu, Muasya & C.H.Stirt.; P. pullata C.H.Stirt.; P. ramulosa C.H.Stirt.; P. rigidula C.H.Stirt.; and P. usitata C.H.Stirt.). Psoraleeae We provide a key for the identification of all ten species along with full descriptions, illustrations, distribution New species maps and some notes on the conservation statuses of the species. South Africa © 2015 SAAB. Published by Elsevier B.V. All rights reserved. Taxonomy

1. Introduction the new world genera are monophyletic, save for some South American species that are presently placed within the genus Otholobium.Onthe The genus Psoralea L. (Leguminosae; sub-family Papilionoideae) other hand, Dludlu et al. (2013) found that the southern African genera belongs to the tribe Psoraleeae, which occurs predominantly in are not monophyletic as the genus Psoralea was nested within the southern Africa, Australia and North America, with outliers in southern African Otholobium. In addition, phylogenetic relationships South America, Mediterranean Europe and Asia (Stirton, 2005). between species could not be established as most of the species formed Psoraleeae sensu Stirton (1981) comprised 6 genera (Psoralea, Hallia polytomies due to low sequence variation in the three DNA regions Thunb., Cullen Medik., Bituminaria Heist ex. Fabr., Orbexilum Raf. and studied. Further molecular phylogenetic studies are in progress to try Otholobium C.H.Stirt. sensu stricto), and ca. 135 species. The Psoraleeae and resolve these issues. tribal study (Stirton, 1981) was preliminary and concentrated mainly The name Psoralea is restricted to taxa sharing the presence of a on the African and Mediterranean genera. Although Stirton (1981) cupulum; a unique cup-shaped structure (resulting from the fusion of recognised that there was more than one genus in the Americas, he intercalary bracts) found at various points along the pedicel (Tucker left all the American species in the genus Orbexilum sensu lato calling and Stirton, 1991). These species have a centre of diversity and ende- for a detailed re-assessment. Key revisionary studies since 1981 have mism in the Greater Cape Floristic Region (GCFR) of South Africa. resulted in the recognition of a number of additional genera among Following extensive fieldwork and studies of herbarium specimens, the New World members of the Psoraleeae. These are as follows: Stirton and Schutte (2012) presently recognize about 70 species in Hoita Rydb., Rupertia J.W. Grimes, Psoralidium Rydb., Pediomelum Psoralea, where up to 30 new species are still only known by manuscript Rydb., Otholobium C.H.Stirt., and Ladeania Egan (Grimes, 1990; Egan names, as depicted in Goldblatt and Manning (2000) and Manning and and Reveal, 2009). In addition, new revisions of Cullen (Grimes, 1997), Goldblatt (2012). The lack of formal descriptions for these species con- Orbexilum (Turner, 2008)andOtholobium (Stirton, 1989) have been founds their identification and impedes assessment of their conserva- done. Thus far, the genus Psoralea has not been revised. Molecular phy- tion statuses. As a result, the necessary conservation action may be logenetic studies by Egan and Crandall (2008) showed that a majority of delayed because before organisms can be conserved they need to be named and described. Therefore, the present study, which is part of an on-going revision of the genus Psoralea, clarifies the taxonomic status fi fl ⁎ Corresponding author. Tel.: +27 21 650 3684; fax: +27 21 650 3301. and provides correct identi cation of the aphyllous (lea ess) Psoralea E-mail address: [email protected] (M.N. Dludlu). species in South Africa, herein referred to as the “P. aphylla complex”.

The P. aphylla complex includes all the members of the genus reduced to unifoliolate leaflets on their seasonal shoots. As mature Psoralea that fall into the broad concept of P. aphylla L., as described plants, they differ in their general habit: P. filifolia is a tall stiff shrub; by Harvey (1862) and generally adopted by subsequent authors. P. fleta is a tall willowy shrub to treelet; while P. gigantea is a large Psoralea aphylla is described as being either leafless, having a subulate tree, with a trunk diameter of up to 40 cm at 1.5 m above the ground. scale instead of a leaf, or sparsely leafy and having a unifoliolate or rarely Psoralea ramulosa, P. rigidula and P. usitata lose their leaves completely asmallfiliform trifoliolate leaf. In the most recent revision of Psoralea, and have glabrous shoots, whereas all the other species have pubescent Forbes (1930) adopted the same description but added the following: shoots. They differ in that P. rigidula is a small, resprouting suffrutex, “leaves unifoliate, 5–17 mm long, linear, acute; present only on with numerous stems branching at the base and is restricted to the Du very young stems, their place in older stems being taken by ovate Toitskloof and Bainskloof Mountains; P. ramulosa is a 2–3 stemmed acute bracts up to 5 mm long”.Basedonthesedefinitions, two resprouting shrub and is endemic to the Cederberg mountains; while other formally published species (P. filifolia Eckl. & Zeyh. and P. the variable, more widespread montane and lowland taxon, P. usitata peratica C.H.Stirt.) would also be included in the complex, making a is a large shrub with 1 –3 stems, terminating in multi-shoot “burst total of three formally published species. Nevertheless, there are sev- branching” with stiff shoots that droop when in flower and has both eral other variants that neither match the type specimens nor the reseeding and resprouting forms. original descriptions of any of the three formally published names, In terms of inflorescence, P. congesta, P. gigantea and P. peratica have and the tendency among botanists and collectors working with pseudo-capitate inflorescences, while all the other seven species have such material has been to lump all leafless Psoralea material under pseudo-spicate inflorescences. A trifid cupulum is found in P. rigidula, the catch-all name “P. aphylla”, hence the “P. aphylla complex”. P. gigantea, P. fleta and P. filifolia, while all the other species have a bifid cupulum. The cupulum overlaps with the calyx in P. aphylla and 2. Materials and methods P. congesta; occurs halfway through the pedicel in P. rigidula, P. filifolia, P. pullata, P. ramulosa and P. usitata; while it is located near the base of Specimens from the Bolus (BOL), British Museum (BM), Compton the pedicel in P. fleta, P. gigantea and P. peratica. The calyx lobes are lon- (NBG), Kew (K), Leningrad (LE), Melbourne (MEL), Missouri (MO), ger than the calyx tube in P. pullata, P. rigidula and P. peratica;thesame Natal (NH), New York (NY), Oxford (OXF), Prague Charles University length as the tube in P. fleta and P. filifolia; whereas in the other species (PRA), Paris (P), Pretoria (PRE), University of KwaZulu-Natal (NU), the lobes are longer than the tube. and Trinity College Dublin (TCD) herbaria were studied. These The existence of such extensive morphological and ecological dis- herbaria hold the largest collections of Psoralea material and thus continuities among members of the P. aphylla complex indicates that capture a significant amount of the diversity in the genus. Additional more than one species can be recognized. Each of the ten species can observations and extensive field collections were done by members be diagnosed using a number of characters as shown in the taxonom- of our research group within the GCFR. Among the characters studied ic treatment. Phylogenetic relationships between species in the were growth form, flower morphology, leaf morphology and pat- genus Psoralea are presently unknown due to lack of resolution in terns of leaf loss and geographic distribution. We also studied some our molecular phylogenetic studies (Dludlu et al., 2013), but studies aspects of seedling morphology including the shape of the first are in progress to address this matter and to determine the mono- eophyll (i.e. the first fully expanded leaves produced by the seedling phyly of the southern African Psoraleeae. Well resolved species in transition to adult leaves), the length of the hypocotyl (i.e. the part level phylogenetic relationships within the genus will shed light on of the stem below the cotyledonary node), and the number of pinnae whether the loss of leaves in Psoralea has occurred as a single evolu- in the metaphyll (i.e. the adult leaves of the seedling). Specimen de- tionary event or has arisen multiple independent times. Details of tails and distribution information are provided as part of the taxo- the description, diagnosis, distribution, conservation status and nomic treatment. These are arranged by Quarter Degree Grid Cells flowering time for each species are provided in the next section. according to Leistner and Morris (1976). Descriptions of vegetation The species are arranged according to their order of appearance in types are based on Mucina and Rutherford (2006) and assessments of the key. the conservation statuses of species were based on the IUCN red list criteria (IUCN, 2012a,b). 4. Taxonomic treatment

3. Results and discussion 4.1. Key to species of the P. aphylla complex

Based on our studies of the morphology of the P. aphylla group, we 1a. Suffrutices, less than 1 m tall; N 5 stems (multi-stemmed) ………… recognize ten distinct species. These species vary in a number of charac- ……….……...... 1. P. rigidula ters including growth form (i.e. herbs, shrubs, trees), persistence strate- 1b. Shrubs or trees, more than 1 m tall; 1–3 stems ……………… gy (i.e. reseeders vs. resprouters), pattern of leaf loss (i.e. leaves lost ………………………………….2 completely, leaves reduced to scales or leaves reduced to a single fili- 2a. Leaves present on flowering shoots, 1–3 foliolate: form leaflet), type of cupulum (i.e. bifidortrifid), positioning of the 3a. Tall, slender, willowy shrubs or small trees; seasonal shoots cupulum on the pedicel, type of inflorescence, pubescence of seasonal emerging on upper parts of tall bare branches; leaves 9–11 mm shoots and geographic distribution. Psoralea aphylla, P. congesta, P. long; inflorescences borne in uppermost axils of seasonal shoots, pullata and P. peratica are all characterised by seasonal shoots with peduncles 1–2 mm long; cupulum trifid, vexillar lobes not bilabi- leaves reduced to scales. They differ in the colour and degree of hairi- ate; standard petal 12–14 mm long; calyx lobes shorter than ness of their seasonal shoots and calyces: lightly covered in black and calyx tube: white hairs (P. aphylla); black hairs covering pedicels and calyces 4a. Plant with slender drooping habit; young shoots with (P. congesta); sparsely covered in white hairs (P. peratica); or densely appressed white pubescence; cupulum trilobed; stan- covered in shaggy black hairs (P. pullata). Geographically, P. aphylla is dard petals purple with darker veins and purple nectar restricted to lower slopes on the Cape Peninsula and the Cape Flats; guides; wing petals white; calyx lobes broadly triangular P. congesta is distributed in the higher reaches of the Cape Peninsula …………………………….2.P. fleta and a few localities across False Bay; P. peratica is a montane species 4b. Plant with distinct tree-like habit when mature; young endemic to the Piketberg Mountains, while P. pullata is a lowland shoots with white densely sericeous pubescence; taxon with a distribution ranging from Hermanus to Potberg Mountain cupulum bilobed with one of the lobes slightly bilabiate; in the east. Psoralea filifolia, P. fleta and P. gigantea have their leaves standard petals mauve, without darker veins and with a M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100 79

small purple ﬂash above the paired callosities; wing 9–11 mm long, purple and white; widespread in Western petals pale mauve; calyx lobes narrowly triangular.....… Cape…………10. P. usitata 3. P. gigantea

3b. Stiff rigid shrubs; seasonal shoots emerging in clusters at end of 4.2. Descriptions of the species stem giving the plant a scruffy or ruffled appearance; leaves 15–20 mm long; inflorescences borne in most axils of seasonal 4.2.1. Psoralea rigidula C.H.Stirt sp. nov. shoots, peduncles 4–5 mm long; cupulum bifid, vexillar lobes Psoralea rigidula C.H.Stirt sp. nov., differs from P. aphylla in its gla- variously bilabiate; standard petal 8–9 mm long; calyx lobes brous multi-stemmed, resprouting suffrutescent habit, up to 0.6 m aris- same length as the calyx tube …..…...... 4. P. filifolia ing from a large, woody, much branched, daucate rootstock; flowers are bright purple to violet, and have a small trifid cupulum, which does not fl ……… 2b. Leaves absent from owering shoots, reduced to scales touch or overlap with the calyx tube. On the other hand, P. aphylla is a ……………………………… ...... 5 sericeous, single stemmed, reseeding shrub to tall tree, 1.5–5.0 m, 5a. Seasonal shoots hairy; calyx tube hairy: with no woody rootstock. It has bluish/mauve flowers and a bifid 6a. Seasonal shoots with clasping and persistent scales; cupulum cupulum, with lobes overlapping or touching the base of the calyx bifid with both lobes non-bilabiate, lobes clasping, situated just tube. Stirton & Schutte in Goldblatt & Manning, Strelitzia 9: 506 below or overlapping with the calyx tube: (Psoralea lucida C.H.Stirt. ined; 2000); Psoralea sp. 10, Stirton and 7a. Tall shrubs to small trees taller than 1.5 m, branching in Schutte (2012). Type: South Africa, Western Cape Province, Worcester upper parts; stems greenish with white lenticels; branches (3319): Limietkop (−CA), 15 Dec 2008, Muasya & Stirton 4333 (BOL!, drooping; inflorescences pseudospicate, laxly flowered, up holo.; BM!, K!, MEL!, MO!, NU!, P!, PRE!, iso.). to 40 flowers per shoot; cupulum just below or overlapping Multi-stemmed suffrutices up to 0.6 m tall, with pinnate leaves in ju- base of calyx at anthesis, lobes deeply cleft, lower pair shal- venile phase, aphyllous in adult phase, resprout after fires from a large lowly cleft; leaf scales acuminate, internodes distant; veins woody daucate rootstock and form dense clumps. Stems erect, numer- of standard purplish in fresh state; lower altitudes of the ous, branching from near woody base, shallowly furrowed; coppice Cape Peninsula and on the Cape Flats ….…………….....5. shoots trifoliolate; leaves 10–15 mm long, 4–5 mm wide; lateral leaflets P. aphylla shorter, excentric, curved, covered on both surfaces with numerous 7b. Short stiff shrubs to 1.5 m tall, branching from near the base; small glands drying reddish; leaves falling off as shoots mature, leaving stems grey; branches stiff, erect, congested at apex of scale-like remnants. Seasonal shoots yellowish-green, emerging from bare stems; inflorescences racemose, up to 5 flowers per base of plant, striate, finely gland-dotted. Scales 1.5 × 1.0 mm, but longer shoot; cupulum not overlapping base of calyx at anthesis, and more attenuated in seedlings, broadly obliquely ovate, glabrous, lobes shallow, lower pair cleft; leaf scales acute, internodes margins minutely ciliate, not tightly congested. Inflorescences borne in congested; veins of standard colourless in fresh state; higher upper axils of seasonal shoots; flowers opening sequentially; pseudo- reaches of the Cape Peninsula and a few localities across spicate; buds glabrous, with lower lobe attenuate and up-curled; pedi- False Bay…………………………………………………….. 6. cel below cupulum 3–4 mm long, 3–4 mm long above cupulum; stiff P. congesta and erect; cupulum trilobed, glabrous, margins ciliate, lobes broadly triangular, densely gland-dotted and drying orange. Flowers 9–12 mm 6b. Seasonal shoots with patent, caducous scales; cupulum bifidwith long, purplish violet and white, bracts reduced to tuft or ring of hairs. – one of the lobes distinctly bilabiate, lobes not clasping, not over- Calyx 6.0 6.5 mm long; lobes subequal, lower lobe slightly longer; lat- lapping with the calyx tube: eral and vexillar calyx lobes acute, straight, triangular; lower lobe 4–5×2–3 mm, acute, broader than the other four lobes; vexillar 8a. Branches rigid, emerging above 1.5 m, with dense calyx lobes fused for up to one third of their length above the tube; branching at each node; seasonal shoots lightly covered calyx shorter than corolla; inner face of calyx lobes finely covered with black and white hairs; scales 3.5–4.0 mm long; inflores- in white hairs with no stubby black hairs; ribs distinctly thickened; cences pseudo-spicate, flowers lax; one flower per axil; pe- glands dense, constant in size, more dense on the tube than the duncles 4–5 mm long; calyx densely covered in long black lobes. Standard petal 8–11 × 9–10 mm, sub-orbicular, purplish violet hairs; Kleinrivier Mountains ………...... ………………………… fading to mauve at edges with purple veins; nectar guides whitish ………………………………………………7. P. pullata with a small central dark purple flash; auricles absent; claw 8b. Branches flexuous, emerging at about 1.0–1.5 m, not densely 4–6 mm long; apex emarginate. Wing petals 10–13 × 4–5mm, branching at each node; seasonal shoots densely covered white, up-curving, fused to, but longer than keel, claw 3–5mm with white hairs; scales 2.5–3.0 mm long; inflorescences long. Keel 10–11 × 3–4mm,claw4–6mmlong.Androecium 10 mm pseudo-capitate, flowers congested; 3–9 flowers per axil; pe- long, scarcely fenestrate. Pistil 8 mm long, stipitate; ovary 1 mm duncles 2 mm long; calyx sparsely covered in short white long, glabrous, sparsely covered in club-shaped glands; style gla- hairs; Piketberg Mountains…………………………………… brous, curved at 4 mm, thickened at point of flexure. Fruits immature, ……………………………………………………… 8. P. peratica papery, reticulate. Seeds 3.5 × 1.5 mm, reinform; hilum central (Fig. 1).

5b. Seasonal shoots glabrous; calyx tube glabrous: 4.2.1.1. Diagnostic features. This distinctive species was previously in- 9a. Plants 1.0–1.5 m tall; stems brownish yellow with white lenticels, cluded within the variation of P. aphylla because of its lack of leaves in resprouting after fires; seasonal growth consists of single shoots its mature shoots. However, it can be distinguished by its glabrous, from all nodes; scales 1.2–2.5 mm long; flowers 17–18 mm multi-stemmed, resprouting, suffrutescent habit, with a maximum long, pale mauve; restricted to Cederberg Mountains.……………. height of 0.6 m. Its flowers are bright purple to violet, and have a 9. P. ramulosa small trifid cupulum, which does not touch or overlap with the calyx 9b. Plants 2–3 m tall; stems greenish grey to pale tan; reseeding and tube. On the other hand, P. aphylla is a sericeous, single stemmed, resprouting after fires; seasonal growth consists of multiple reseeding shrub to tall tree, with a height of 1.5–5.0 m. It has bluish/ shoots from terminal nodes; scales 2.5–3.5 mm long; flowers mauve flowers and a bifid cupulum, with lobes usually overlapping or 80 M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100

a b c d

p h

l n o k

Fig. 1. Vegetative and reproductive morphology of P. rigidula:(a)flower bract; (b) cupulum opened up; (c) flower bud; (d) fruit; (e) cupulum; (f) mature flower; (g) standard petal; (h) standard petal folded; (i) wing petal; (j) keel petal; (k) calyx opened up: upper lobes to the left; (l) pistil; (m) general habit; (n) seed; (o) close-up of shoot; (p) seasonal shoot in flower. Voucher: Esterhuysen 35742 (BOL). Scale bars: a–l, n, 1 mm; m, 10 cm; p, 2 cm; 0, 10 mm. Artist: A. Beaumont. touching the base of the calyx tube. Specimens of this species collected above sea level (Fig. 2). It occurs in seasonal seepages in Hawequas by Ms Elsie Esterhuysen (28213, 33766) from the NE ridge of Du Toit's Sandstone Fynbos i.e. FFs 10 (Rebelo et al., 2006). Flowering time is Peak above Delabat Ravine were annotated previously in some herbaria November to April. under the manuscript name of “P. lucida”. These collections, which may well represent a distinct species, are here included within P. rigidula 4.2.1.3. Etymology. The specificepithetrigidula is derived from the Latin pending further investigation. They have coarser and thicker stems word rigidulus (adj.) meaning stiff or rigid, and refers to the distinctive and shoots (much like Indigofera gifbergensis C.H.Stirt.), larger flowers, rigid dwarf habit of the plant. 15–17 mm long and very short pedicels, 1–2 mm long, cupulum almost at the base of the pedicel, with dense coarsely glandular scales and 4.2.1.4. Conservation status Psoralea. rigidula has a very narrow distribu- calyces, and (fide Esterhuysen) blue flowers. tion range and is not abundant, except in the few large colonies where it occurs. Recruitment must occur at a slow rate and is evidenced by the 4.2.1.2. Distribution and habitat. A narrow endemic restricted to the variable age class distribution of plants in the population, ranging mountains around Bainskloof and Du Toit's Kloof, at 500–1500 m from small few-stemmed rootstocks to very large ones producing scores M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100 81

Fig. 2. Known distribution of P. rigidula: section of map shown is the south-western tip of South Africa. of stems. Based on the IUCN red list criteria (IUCN, 2012a,b) we classify (9–11 mm long) on some shoots. Scales 3.0–5.0 × 0.7–0.9 mm, smaller the taxon as VU B1b(i,ii). on flowering shoots, longer than petioles, not fused to petioles, obliquely narrowly ovate to subulate, clasping, glabrous and not tightly 4.2.1.5. Additional specimens examined. 3319 (Worcester): above Delabat congested. Inflorescences borne in uppermost axils of seasonal shoots, Ravine, NE Du Toits Kloof (−CA), 28 Feb 1959, Esterhuysen 28213 pseudo-spicate, congested, up to three flowers per axil; pedicel below (BOL); observation point, Bainskloof Mountains (−CA), 23 Jan 1982, cupulum 1–2 mm long, stout and rigid, shorter than the subtending Esterhuysen 35742 (BOL, E, NBG, PRE); Bainskloof Pass (−CA), 5 Mar leaflet; cupulum trilobed with one of the lobes scarcely developed, 1953, Levyns 9898 (BOL); 30 Nov 2007, Muasya & Stirton 3390 (BOL); broadly triangular, glabrous but hairy on lobes; pedicel above cupulum 15 Mar 2008, Muasya & Stirton 3882 (BOL, K, MO); 13 Sep 2008, Muasya 3–4 (5) mm long. Flowers 12–16 mm long, light purple, maturing se- & Stirton 3958, 3962 (BOL, NBG, PRE); 4 Oct 2008, Dludlu & Stirton 30 quentially; bracts reduced to tuft or ring of hairs. Calyx 7–8 mm long, (BOL, NBG, PRE); 22 Jun 2010, Muasya & Stirton 5343 (BOL); bottom tube 2.0–4.5 mm long, glabrous externally, inner face of calyx lobes of hill, Worcester side of Bainskloof (−CA), 4 Oct 2008, Dludlu, Muasya & sparsely covered in small black stubby hairs; lobes shorter than the Stirton 28 (BOL, NBG, PRE); 1.5 km south of Hugenootkop, Klein tube, unequal; lateral and vexillar calyx lobes acute, straight, triangular; Drakenstein mountains (−CC), 14 Jan 2007, Helme 4511 (NBG); NE lower lobe 6 mm long, 2.0–2.5 mm wide, acuminate and narrower than ridge of Du Toit's Peak, above Delabat Ravine (−CC), 7 Feb 1975, the other four lobes; vexillar calyx lobes free above the tube; calyx shorter Esterhuysen 33766 (BOL); Haelhoek Spitzkop (−CC), 2 Jan 1947, than corolla; ribs distinctly thickened; glands dense, constant in size and Esterhuysen 14550 (BM, BOL, K); slopes of Winterberg facing Haelhoek, equally distributed across tube and lobes; lobe margins ciliate. Standard Sneeukop (−CC), 15 Mar 1957, Esterhuysen 28229 (BM, BOL, K); petal 12–14 × 12–13 mm, orbicular; auricles present; apex rounded or between Haelhoek Sneeukop and Winterberg, above stream flowing obtuse; claw 4 mm long, elongated and narrow. Wing petals into Wemmershoek Valley (−CC), 14 Mar 1959, Esterhuysen 28233a 12.5–15.0 × 4.5–5.5 mm, up-curving, auricles present, fused to but longer (BM, BOL). than keel, claw 4 mm long. Keel 11–13 × 4 mm, claw 5–7mmlong.An- droecium 10 mm long, scarcely fenestrate. Pistil 8 mm long, stipitate; 4.2.2. Psoralea fleta C.H.Stirt sp. nov. ovary 1 mm long, glabrous, sparsely covered in club-shaped glands; Psoralea fleta C.H.Stirt sp. nov., differs from P. gigantea in its more style glabrous, curved at 4 mm, thickened at point of flexure. Fruits slender drooping habit, young shoots with appressed white pubescence, 5.0 × 2.5–3.0 mm, finely reticulate, papery. Seeds 4.0 × 2.5 mm, reinform; trilobed cupulum, purple standards with darker veins and purple nectar hilum sub-central, blackish grey (Fig. 3). guides, white wing petals, and broadly triangular calyx lobes. Mean- while, P. gigantea has a distinct tree habit, densely sericeous young shoots, a bifid cupulum, with one of the lobes slightly bilabiate, mauve 4.2.2.1. Diagnostic characters. Psoralea fleta differs from the rest of the standard petals with a small purple flash above the callosities, pale P. aphylla group by its filiform leaflets (9–11 mm long) borne on its mauve wing petals and narrowly triangular calyx lobes. Stirton and flowering shoots. In habit it ranges from tall willowy shrubs to small Schutte (2000); Psoralea sp. 4, Stirton and Schutte (2012). Type: South slender trees, with drooping branches when in full flower producing a Africa, Western Cape, Worcester (3319): Bainskloof Pass (−CA), 22 mass of intensely sweet-smelling flowers. Each flower is subtended by Oct 1986, Stirton & Snijman 11226 (NBG!, holo.; BM!, K!, NBG!, P!, iso.). atrifidcupulum.Psoralea fleta is similar to P. gigantea in that they Tall, slender, willowy shrubs or small trees, 4–5mtall,reseeder,usu- both have seasonal shoots emerging on upper parts of tall bare ally dominant and forming dense stands. Stems single, erect, woody branches, filiform, unifoliolate leaves in seasonal shoots, inflorescences throughout and may be weakly furrowed, but generally rounded; borne in the uppermost axils of seasonal shoots and calyx lobes that branches pale green, lightly gland-dotted, flexuous, emerging at upper are shorter than the calyx tube. However, P. gigantea can be separated parts of tall bare stems. Seasonal shoots drooping, especially when by its distinct tree habit when mature, densely sericeous hairy young in flower, covered in a close white pubescence, smooth, striate and shoots, bilobed cupulum with one of the lobes slightly bilabiate, non-pustulate; leaves absent, but may have a tiny filiform leaflet mauve standard petals without darker veins and with just a small 82 M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100

b a

d e

l i

h jk

Fig. 3. Vegetative and reproductive morphology of P. fleta: (a) cupulum; (b) pistil; (c) mature flower; (d) standard petal; (e) standard petal folded; (f) wing petal; (g) keel petal; (h) calyx opened up: upper lobes to the left; (i) close-up of shoot; (j) unifoliolate leaf; (k) close-up of scale; (l) seasonal shoot in flower. Voucher: Stirton & Snijman 11226 (NBG). Scale bars: a–h, j, k, 1 mm; i, l, 10 mm. Artist: A. Beaumont.

purple flash above the paired callosities, pale mauve wing petals, and fires it germinates rapidly and once the seedling reaches about 10 cm narrowly triangular calyx lobes. tall it forms numerous basal lateral branches. These leafy mounds are densely pinnately-leaved. After a few weeks the basal branches have 4.2.2.2. Distribution and habitat. This species is restricted to Hawequas all abscised leaving a single shoot which extends rapidly reaching Sandstone Fynbos i.e. FFs 10 (Rebelo et al., 2006). It occurs on the almost 3 m in the first season, gradually producing fewer and fewer mountains between Wellington, Ceres and Tulbagh (Limietberge, leaflets until only scales are produced by the end of the first season. Elandskloofberge, Witsenberge), with a wide altitudinal range from (250) 600–2050 m above sea level but is generally found at higher alti- 4.2.2.3. Etymology. The name fleta is derived from the Latin word fletus tudes in wet seepages and ravines on eastern slopes (Fig. 4). Flowering (m) — weeping or tears, and refers to the masses of pendulous flowers time is September to March. This is a very fast growing species. After which glisten like tears when viewed against the sun, especially after rain. M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100 83

4.2.2.4. Conservation status. Although P. fleta has a wide distribution it Cape Town (3318): Jonkershoek Nature Reserve, Stellenbosch (−DD), is only locally abundant. Based on the IUCN red list criteria (IUCN, 29 Oct 2008, Dludlu, Muasya & Stirton 57 (BOL!, holo.; K!, MO!, NBG!, 2012a,b) we classify the taxon as VU B2b(i,ii). PRE!, iso.). Trees, 6–8 m tall, reseeder, forming dense clumps especially in deep 4.2.2.5. Additional specimens examined. 3319 (Worcester): Saron, fire-protected valley sites. Stems erect, reaching a trunk diameter up to Tulbagh (−AA), Nov 1941, Stokoe s.n. (PRE); Elandsfontein farm, 40 cm, grey, woody throughout, not furrowed, start branching in below Elandskloofberge (−AA), Stirton 11241 (BM, K); Mitchells Pass upper portions above 2 m; young plants densely branched and compact, (−AD), 1873, Bolus 2607 (K); 5 May 1941, Walgate 379 (PRE); 6 Jan broom-like, very fast growing; branches robust, patent, with two or 1941, Esterhuysen 6148 (BOL); bottom of Mitchell's Pass, Tulbagh side three branches at each node, culminating in drooping shoots when (−AD), 30 Nov 2007, Muasya & Stirton 3341, 3342 (BOL); Ceres older. Seasonal shoots densely sericeous when young, becoming sparsely (−AD), 2 Oct 1933, Levyns 4663 (BOL); Bainskloof (−CA), 28 Oct hairy, mostly within grooves of the leaflets, striate, drooping, dark 1928, Hutchinson 1065 (BM, BOL, K); 5 Mar 1953, Levyns 9897 (BOL); green, smooth, lightly gland dotted; glands drying black; leaves 1 (3)- 23 Oct 1966, Taylor 6987a (PRE); 21 Jan 1979, Grobbelaar 2204 (PRE); foliolate on seasonal shoots, though most flowering shoots have a Meyer 1412 (PRE); 30 Nov 2007, Muasya & Stirton 3385 (BOL); 15 Mar filiform leaflet up to 30 mm long, glabrous. Scales 1.5 × 1.2 mm, nar- 2008, Muasya & Stirton 3883 (BOL, K, MO, NBG); 13 Sep 2008, Muasya rowly triangular, erect to spreading, glabrous, not tightly congested. & Stirton 3959, 3960 (BOL); 4 Oct 2008, Dludlu, Muasya & Stirton 29 Inflorescences of 2–4 flowers, borne in uppermost axils of seasonal (BOL); 22 Jun 2010, Muasya & Stirton 5345 (BOL); Du Toits Kloof Pass shoots, pseudo-capitate, maturing sequentially; pedicel below Tunnel (−CA), 28 Sep 1981, Stirton 9478 (PRE); along path from cupulum 1–2mmlong,filiform and flexuous; pedicel above cupulum Bainskloof Hotel to Baviaanskloof (−CA), 14 Nov 1965, Taylor 6604 3–4 mm long; cupulum bilobed with one of the vexillar lobes vari- (PRE); Baviaanskloof (−CA), 14 Nov 1965, Taylor 6603 (PRE); ously bilabiate, glabrous, but ciliate along lobe margins; lobes equal- Bainsberg, Bobbejaansrivier path on NW facing slope (−CA), 28 Dec ly developed, glandular and narrowly triangular. Flowers 10–14 mm 1989, Bean 2129 (BOL, K); Wolvekloof Forest Reserve (−CA), 28 Oct long, dark mauve, with a small purple spot above the white swollen 1946, Barker 4244 (BOL, NBG); Brandwagt (−CB), 14 Mar 1949, Van appendages at base of blade. Calyx 10 mm long, tube 3.5 mm long, Breda 347 (PRE); Zachariashoek, Kasteelkloof ( −CC), 24 Mar 1983, glabrous, tube shorter than the lobes; calyx lobes unequal, lower Viviers 1140 (PRE); Berg River Hoek, Paarl District (−DA), 8 Apr 1944, lobe much longer than the other four lobes; lateral and vexillar Compton 15637 (NBG); west side of Keeromsberg (−DA), 7 Nov 1943, calyx lobes acuminate, falcate, and narrowly triangular; lower lobe Esterhuysen 9204 (BOL, PRE). 8.0 × 2.2 mm, broader than the other four lobes; vexillar calyx lobes fused for up to one third of their length above the tube; calyx 4.2.3. Psoralea gigantea Dludlu, Muasya & C.H.Stirt sp. nov. shorter than the corolla; inner face of calyx lobes densely covered Psoralea gigantea Dludlu, Muasya & C.H.Stirt sp. nov., differs from in small black stubby hairs; ribs distinctly thickened; glands dense, P. fleta in its distinct tree habit when mature, with densely sericeous constant in size, glands equally dense on lobes and the tube. Standard young shoot tips, bilobed cupulum with one of the lobes slightly bilabi- petal 10 × 9 mm, sub-orbicular; auricles well-developed; apex emarginate, mauve standard petals without darker veins and a small purple ate; claw 5 mm long, elongated and narrow. Wing petals 12–14 × 4 mm, flash above the paired callosities, pale mauve wing petals, and narrowly up-curving and longer than keel, fused to but longer than keel; claw 3– triangular calyx lobes. Psoralea fleta is a slender willowy shrub to small 6mmlong.Keel 11–12 × 3–4 mm; claw 5–7mmlong.Androecium tree, with appressed white pubescence on young shoot tips, trilobed 10 mm long, upper end curved inwards, 10th stamen lightly attached, cupulum, purple standard petals with darker veins and purple nectar fenestrate. Pistil 11 mm long; ovary 2 mm long, with very few small guides, white wing petals, and broadly triangular calyx lobes; Psoralea stalked glands; style 6 mm long, swollen beyond point of flexure, sp. 6, Stirton and Schutte (2012). Type: South Africa, Western Cape, curved at 4 mm. Fruits 6 × 3 mm, coarsely reticulate, membranous,

Fig. 4. Known distribution of P. ﬂeta: section of map shown is the south-western tip of South Africa. 84 M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100

h c

k e

f l

Fig. 5. Vegetative and reproductive morphology of P. gigantea:(a)matureflower (front-view); (b) flower bud; (c) cupulum; (d) unifoliolate leaf; (e) calyx opened up: upper lobes to the left; (f) seasonal shoot in flower; (g) mature flower (side-view); (h) standard petal; (i) wing petal; (j) keel petal; (k) pistil; (l) androecium. Voucher: Dludlu, Muasya & Stirton 57 (BOL). Scale bars: a–e, g–l, 1 mm; f, 4 cm. Artist: A. Bello. black; apex beaked. Seeds 4.0 × 2.5 mm, reniform; hilum sub-central, veins and purple nectar guides, white wing petals, and broadly triangu- blackish grey (Fig. 5). lar calyx lobes.

4.2.3.2. Distribution and habitat. The species is restricted to protected 4.2.3.1. Diagnostic characters. Psoralea gigantea is the only member of the valleys from Jonkershoek to Mitchell's Pass (Fig. 6), at altitudes P. aphylla complex which grows into a large tree. Its flowering shoots 300–700 m above sea level. It inhabits moist habitats such as river have filiform unifoliolate leaves (up to 30 mm long) with 2–4 flowers and stream margins in Kogelberg Sandstone Fynbos i.e. FFs 11 (Rebelo per axil. Unlike P. fleta the flowers are dark mauve and only faintly et al., 2006). Flowering time is September to December. fragrant. Psoralea fleta resembles P. gigantea in having filiform unifoliolate leaves in its flowering shoots. It differs in its more slender drooping 4.2.3.3. Etymology. The specificepithetgigantea is derived from the Latin habit, young shoots with appressed white pubescence, trilobed word gigantes (m.), meaning large or gigantic, and alludes to the very cupulum, intensely fragrant flowers, purple standard petals with darker large size of this species — one of only three true trees in the genus. M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100 85

Fig. 6. Known distribution of P. gigantea: section of map shown is the south-western tip of South Africa.

The others are P. arborea Sims and an undescribed species from the Ecklon & Zeyher 1532 (S! (S08-12030), lecto.; GOET!, L!, M!, MO!, S! Swartberg and Baviaanskloof mountains (Psoralea sp. 25; Stirton and (S08-12028), iso.). Lectotype chosen here. Schutte, 2012). Large shrub, 3.5–4.0 m tall, reseeder, forms dense clumps. Stems single, woody throughout, terete, not furrowed, greyish, with white lenti- 4.2.3.4. Conservation status. Psoralea gigantea is only known from a few cels; branches stiff, emerging at height of about 1.5 m, several localities and is not abundant. However, it occurs in protected areas. branches produced per node, giving the plant a somewhat ruffled or Therefore, based on the IUCN red list criteria (IUCN, 2012a,b) we classify scruffy appearance. Seasonal shoots dark green, smooth, 1–3-foliolate; the taxon as VU B1b(i,ii). leaves 15–20 mm long, hemispherical in cross-section with a straight, sharp and hard mucro at the apex; leaflet glands in dry state visible 4.2.3.5. Additional specimens examined. 3318 (Cape Town): Jonkershoek with a ×10 hand lens, reddish black, impressed on surface, denser on Nature Reserve (−DD), 5 Oct 2008, Dludlu, Muasya & Stirton 35 (BOL); upper surface; petioles 2 mm long. Scales 3.5–4.0 × 1.0 mm, persistent, 16 Nov 2008, Muasya & Stirton 3201b, 3203 (BOL); 21 Oct 1970, glabrescent, not tightly congested. Inflorescences borne in most axils of Grobbelaar 1162 (PRE); 23 Oct 1963, Taylor 5487 (NBG); 1 Oct 1958, seasonal shoots, pseudo-spicate, congested (up to 40 flowers per Werdermann & Oberdieck 374 (BOL, PRE); Swartboskloof, Stellenbosch shoot), with 1–3 flowers per axil; pedicels below cupulum 4 mm long, (−DD), 17 Oct 1960, Van Rensburg 2124 (PRE); south of Jakalsvlei stout and rigid; cupulum bilobed with one of the vexillar lobes variously Rivier, Jonkershoek Nature Reserve Catchment (−DD), 2010, Dlodlo & bilabiate; pedicel above cupulum 4 mm long; cupulum lobes equally de- Stirton 34 (BOL); Coetzenburg, above dam (−DD), Walters s.n. (PRE). veloped, narrowly triangular, hairy on lobe margins and along adaxial 3319 (Worcester): Mitchell's Pass (−AD), 5 Apr 1974, Goldblatt midrib; cupulum glands dense, conspicuous, reddish orange. Flowers 1346 (PRE); Haelhoek Sneeukop foothills (−AD), Paarl Division, Stokoe 16–20 mm long, mauve, maturing sequentially. Calyx tube 3–4mm s.n (SAM 55824); Mitchell's Pass at junction of R46 & R43 (−AD), 5 Oct long, completely glabrous on the outside, but inner face of calyx lobes 2008, Dludlu, Muasya & Stirton 35 (BOL); Krom Riviers Kloof, off Du sparsely covered in small black stubby hairs; calyx lobes subequal, Toitskloof (−CA), Esterhuysen 28276 (BOL); Donkerkloof, Paarl same length as the calyx tube; lateral and vexillar calyx lobes acute, (−CC), Esterhuysen 7738 (BOL); Haelhoek Sneeukop (−CC), 14 Mar straight, lanceolate; lower lobe 4 mm long, 2 mm wide, acute, broader 1959, Esterhuysen 28232 (BOL, NBG); Tierkloof, Wemmershoek than the other four lobes and slightly longer; calyx shorter than corolla; (−CC), Gray s.n. (BOL); Boesmanskloof, seep above Bobbejaans glands dense, small, constant in size and equally distributed across the River, Boland hiking trail (−CC), 11 Dec 2000, Low 15783 (BOL); calyx tube and lobes. Standard petal 8–9×10 –12 mm, broadly ovate; Wemmershoek Mountains, north of dam, Veilingskloof (−CC), 21 Dec claw 5 mm long, narrow; apex emarginate; auricles well developed. 2004, Helme 3305 (NBG); eastern slopes of Franschoek Pass (−CC), 3 Wing petals 12–14 × 4 mm, cultrate with a strong fold along the midline, Dec 1981, Stirton 10120 (PRE). up-curving, fused to but longer than keel; auricles present, swollen; 3420 (Montagu): Marloth Nature Reserve (−AB), 5 Oct 2008, claw 4–5mmlong.Keel 10–11 × 4–5 mm; claw 6 mm long; apex round- Dludlu, Muasya, Chimphango & Stirton 37 (BOL). ed. Androecium 9 mm long, fenestrate. Pistil 9mmlong,pedicellate; 3418 (Simonstown): Lourensford Estate (−BB), 11 Oct 1944, Parker ovary 1 mm long; style filiform, 7 mm to point of flexure, swollen before 3935 (BOL). point of flexure, length of curvature 3 mm; stigma penicillate. Fruits 3419 (Caledon): Van der Stel (−AA), 12 Nov 1929, Smith 4854 5 × 3 mm, papery, strongly reticulate, black. Seeds 2 × 1 mm, black; (PRE); west of Viljoens Pass, Caledon Mountains (−AA), Stokoe 7885 hilum sub-central, aril well-developed, 1 mm long (Fig. 7). (BOL). 4.2.4.1. Diagnostic characters. This species is similar to P. fleta and 4.2.4. Psoralea filifolia Eckl. & Zeyh. P. gigantea in that it has unifoliolate leaves in its seasonal shoots. How- Enum. Pl. Afric. Austral. 2: 227. (1836); Stirton and Schutte (2000, ever, P. filifolia may also have trifoliolate leaves on younger stems in 2012). Type: South Africa, Western Cape, Worcester (3319): “in which the terminal leaflet is twice the length of the lateral leaflets. In humidis (Altit. II) fruticem prope Waterfall in valle Tulbagh” (−AA), addition, P. filifolia is a large shrub with an untidy appearance due to 86 M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100

c ab

j i m p

o l n

Fig. 7. Vegetative and reproductive morphology of P. filifolia:(a)flower bract opened up; (b) flower bracts; (c) flower bud; (d) cupulum; (e) mature flower; (f) standard petal; (g) standard petal folded; (h) wing petal; (i) keel petal; (j) stigma; (k) pistil; (l) calyx opened up: upper lobes to the left; (m) seasonal shoot in flower; (n) close-up of stipules; (o) leaflets; (p) general habit; (q) scales. Voucher: Stirton 11199 (K, PRE). Scale bars: a–l, n, o, q, 1 mm; m, 10 mm; p, 25 cm. Artist: A. Beaumont. its burst branching, while P. fleta and P. gigantea do not burst branch, on shale and in seepages in sandveld, at altitude 20–140 m above sea branching only in upper parts of tall stems. Its stem has white level. This species is found in Breede Shale Renosterveld i.e. FRs 8, lenticels, a feature which is absent in the other two species. Hopefield Sand Fynbos i.e. FFd 3, Atlantis Sand Fynbos i.e. FFd 4 and Cape Flats Sand Fynbos i.e. FFd 5 (Rebelo et al., 2006). Flowering time 4.2.4.2. Distribution and habitat. This species occurs on stream banks and is September to March. seepages in the Verlorenvlei system in the northern part of its range where it is abundant while southwards and eastwards it is now less fre- 4.2.4.3. Etymology. The specific epithet filifolia is a compound word quent (pers. comm., Louw 2010). Despite a number of thorough derived from the Latin words filli- (fine) and folius (n. leaf) and refers searches on the Cape Flats and in the Tulbagh area we were unable to to the fine or thread like leaves. find extant populations and it is probably now extinct in these areas. There is a healthy population in the Rondeberg Private Nature Reserve 4.2.4.4. Conservation status. The Red List of South African Plants (version (Fig. 8). It occurs along stream banks and seepages in lowland fynbos 2014.1; Raimondo et al., 2009) lists it as EN B1ab(ii,iii,iv,v). It occurs in a M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100 87

Fig. 8. Known distribution of P. ﬁlifolia (squares) and P. aphylla (circles): section of map shown is the south-western tip of South Africa.

few protected areas and is not usually very abundant. Stirton and in S. Afr. J. Bot 64: 244 (1998); Stirton and Schutte (2000); Stirton and Raimondo (2008) reported that it had largely disappeared from its Schutte (2012); non Reichenbach (1822). Type: South Africa, Western lowland seepage habitats and that most of this species' habitat has Cape, Cape Town (3318): Cape of Good Hope, Oldenland 685 (G!, lecto.; been transformed for the cultivation of vineyards, wheat and potatoes, designated by Stirton in Turland & Jarvis (Ed.), Taxon 46: 479 (1997)). and that the decline continues. [Genista spartium coeruleum capitis bonae spei. Brey., Exot. Pl. cent. 66, t.25 (1678)]. 4.2.4.5. Additional specimens examined. 3218 (Clanwilliam): Verlorenvlei, Psoralea decidua Berg., Descr. Pl. Cap. 220 (1776) nom. illegit. non north of Grootdrif, West coast near Elands Bay (−AD), 23 Sep 2002, Thunb. Low 7555 (BOL). Psoralea denudata Hoffmannsegg, Verzeichn. Pflanz. 190 (1823), 3318 (Cape Town): Rondeberg Private Nature Reserve (−AD), 6 Dec nom. superfl. 2008, Muasya & Stirton 4321 (BOL); Mud River (−AD), 21 Sep 1981, Psoralea jacquiniana Eckl. & Zeyh., Enum. 226 (1836) nom. illegit., Stirton 9361 (PRE); de Wet s.n. (NBG); Groote Post farm near Darling– non.DC.nec G. Don. Mamre Road (−AD), 17 Oct 1986, Stirton 11199 (K, PRE); near Cape Tall shrubs or small trees, 1.5–5.0 m tall, erect, often forming dense Town (−CD), Harvey s.n. (TCD). colonies, reseeder. Stems single, rarely more, woody throughout, terete, 3319 (Worcester): Vierentwintigrivier (− AA), Drège s.n. brown when older, lenticelled; branches erect, becoming arcuate as (P03050175); Tulbagh Waterfall (−AA), Stokoe s.n. (MO, OXF, PRA); they lengthen, yellowish green with white lenticels, commencing at a Tulbagh (−AA) Ecklon & Zeyher 1532 (MO, OXF, PRA, SAM); Steendal, height of about 1 m, flexuous, minutely glandular. Seasonal shoots near Tulbagh (−AA), Hutton s.n. (TCD); Steenboksberg (−AA), 23 Oct densely covered in white to silvery pubescence, with a few randomly 1966, Taylor 6987a (STE); Saron, Tulbagh Kloof (−BB), 14 Feb 1922, scattered glands; leaves absent in mature plants, but seedlings are Pole Evans 485 (PRE); Malbrokskloof (−CA), Drège s.n. (P). 1–3-foliolate; some side branches of young stems may have 3-foliolate 3418 (Simonstown): south of Pringle Bay (−BD), 12 Sep 1969, (15–30 mm long) leaves otherwise, seasonal shoots covered in scales. Boucher 627 (PRE); Malbrokskloof (−CA), Drège s.n. (P). Scales 4.0–6.0 × 1.5–3.0 mm, green, persistent, becoming woody, lance- Without precise locality: Anonymous s.n. (P 03496091); Anon. s.n., ex olate, clasping or free on younger shoots, glabrous, with few hairs inside. Hort. Kew (BM); Drège s.n. (P 03496119); Sonnerat s.n. (P 03496123); Inflorescences borne in upper axils of seasonal shoots, flowers opening Viellard & Deplanche s.n. (P 03368446); Cape Peninsula, Nelson s.n. sequentially, pseudo-spicate, lax, 10–30 flowers per shoot; buds hairy; (BM, K). pedicels 3–4 mm long below cupulum, stout, erect, rigid, hairy; cupulum deeply bilobed, vexillar lobes almost fused to give a small 4.2.5. Psoralea aphylla L. cleft, lobes equally developed and usually overlapping or touching the Pl. Rar. Afr. 15 (1760); Amoen. Acad. 6: 93 (1763); Houttyn, Handl. base of the calyx tube, broadly subulate, cuspidate, gland dotted; lobes Pl. Kruidk. 5: 552 (1776); Poir. in Lam. Encycl. 5: 681 (1804); Dietrich, glabrous with ciliate margins; tube hairy, minutely glandular; pedicels Lex. Gart. Bot. 7: 607 (1807); Sims in Bot. Mag. 42: t.1727 (1815); above cupulum 2–4 mm long, hairy. Flowers 11–16 mm long, bluish Loddiges, Bot. Cab. 3: t. 221 (1818); DC., Prodr. 2: 217 (1824) pro mauve/purple and white with darker purple veins, maturing more parte; E. Mey. in Linnea 7:165 (1832); G. Don., Gen. Syst. 2: 202 or less simultaneously; flower bracts inconspicuous, replaced by a (1832); Eckl. & Zeyh., Enum. 227 (1836); E. Mey., Comm. 84 (1936); tuft or ring of hairs. Calyx shorter than corolla; lobes shorter Presl, Bot. Berm. 53 (1844); Klinsman, Clav. Breyn. 9 (1855); Harv. in than calyx tube, sub-equal, lower lobe slightly longer; lower lobe Harv. & Sond., Fl. Cap. 2: 145 (1862); Hamer, Wild Flowers Cape 23, t. 3.0–3.5 × 1.0–2.5 mm, acute, broader than the other four lobes; vexillar 71 (1926); Salter in Adamson & Salter, Fl. Cape Penins. 489 (1950); lobes free above the tube; tube 2.5–3.0 mm long, lightly covered in black Forbes in Bothalia 3: 5 (1930); Kidd, Wild Flowers Cape Penins. T. and white hairs; inner face of calyx lobes sparsely covered in small black (1973); Burman & Bean, South Afr. Wild Fl. Guide 5: 116 (1985); Stirton stubby hairs; calyx glands raised, constant in size, denser on tube; 88 M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100

h a

c j d

e l

Fig. 9. Vegetative and reproductive morphology of P. aphylla: (a) mature flower (side-view); (b) flower bud; (c) scale; (d) cupulum; (e) calyx opened up: upper lobes to the left; (f) seasonal shoot in flower, (g) mature flower (front-view); (h) standard petal; (i) wing petal; (j) keel petal; (k) pistil; (l) androecium. Voucher: Muasya & Stirton 3400 (BOL). Scale bars: a–e, g–l, 1 mm; f, 2 cm. Artist: A. Bello.

accrescent in fruit. Standard petal 8–10 × 9–11 mm, sub-orbicular, blu- papery, reticulate, glabrous. Seeds 3.0 × 2.0–2.3 mm, 1.5 mm thick, ish mauve to purple with darker purple veins, with two free vertical reniform, blackish brown; hilum rim flanged, sub-terminal (Fig. 9). 2 mm long white appendages 2 mm above claw with a purple flash in between; apex emarginate; claw elongated and narrow, 3–5mmlong; 4.2.5.1. Diagnostic characters. Psoralea aphylla differs from all other auricles well-developed; Wing petals 10–13 × 3–5 mm, white, longer members of the complex by the presence of large, persistent and mostly than keel; claw 3.0–4.5 mm long. Keel 8.0–12.0 × 2.0–3.5 mm; claw clasping scales on its seasonal shoots. Its flowers have a bifidcupulum 4–6mmlong.Androecium 9–10 mm long. Pistil 9 mm long, ovary (lower pair with a small cleft) that clasps up to the base of the calyx 1.5 mm long; style filiform, 7 mm to point of flexure, glabrous but (can be free in fruiting phase) and is covered in white or silvery hairs. with stalked club-shaped glands on upper margin, pedicellate, length Psoralea rigudula, which is often identified as P. aphylla in herbarium of curvature 4 mm; stigma penicillate. Fruits 4.0 × 2.5 mm, black, specimens due to its lack of leaves is a resprouting, suffrutex with M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100 89

g c

i d

e f k

Fig. 10. Vegetative and reproductive morphology of P. congesta:(a)matureflower (front-view); (b) mature flower (side-view); (c) scale; (d) cupulum; (e) calyx opened up: upper lobes to the right; (f) seasonal shoot in flower; (g) standard petal; (h) pistil; (i) androecium; (j) keel petal; (k) wing petal. Voucher: Dludlu & Stirton 39 (BOL). Scale bars: a–e, g–k, 1 mm; f, 3 cm. Artist: A. Bello. multiple stems, reaching a maximum height of 0.6 m. Its stems are glan- there. Its altitude ranges from 10 to 300 m above sea level. Growing on dular and without lenticels. Its seasonal shoots are erect even when in Peninsula Sandstone Fynbos i.e. FFs 9 and Cape Flats Sand Fynbos i.e. FFd flower and the flowers are purple-violet. In contrast, P. aphylla is a 5(Rebelo et al., 2006); open areas, stream banks, gulleys and seepages. reseeding, tall willowy shrub or small tree that grows up to 5 m tall. It Flowering takes place from September to April. has greenish-tan stems with storied white lenticels, seasonal shoots that become floppy when in flower, and the flowers are bluish mauve. 4.2.5.3. Etymology. The specificepithetaphylla is a compound Greek Psoralea aphylla is most similar to P. congesta and P. usitata in that all word meaning ‘without leaves’ and is derived from the Greek words three species have their leaves reduced to persistent scales in the seasonal ά- (a-, without, not) and φύλλο (leaf). shoots. Psoralea congesta differs from P. aphylla in its mostly leafless erect stiff grey stems bearing many short compact broom-like short seasonal 4.2.5.4. Conservation status. The Red List of South African Plants (version shoots aggregated at the apex, black-haired pedicels, congested inter- 2014.1; Raimondo et al., 2009) lists it as Least Concern (LC). This taxon nodes and appressed scales, and black-haired calyces. Psoralea usitata is was not selected in any one of their four screening processes for an erect reseeding or resprouting shrub without congested scales, a highlighting potential taxa of conservation concern for detailed assess- cupulum that has two lobes, one with a deeper cleft (making it appear tri- ment and was hence given an automated status of LC. However, as fid). The cupulum never overlaps with the calyx, it is situated about half- P. aphylla has been broken up into a number of distinct taxa (this way along the pedicel. Furthermore, in P. aphylla the seasonal shoots are paper), it is seriously endangered because of habitat conversion for covered with white silvery hairs, while in P. usitata they are glabrous. human settlement. It used to be quite common along the lower slopes The distribution ranges of the two are allopatric. of the Cape Peninsula and in seepage areas across the Cape Flats. Today it is mostly restricted to a few areas of the lower protected 4.2.5.2. Distribution and habitat. Psoralea aphylla is scattered across the bases of mountain streams and gulleys as they abut on human settle- Cape Peninsula and surrounding low foothills of mountains (Fig. 8). Al- ments. It is extinct in most of the areas from which it was formerly though it used to be widespread on the Cape Flats, it is now quite scarce known on the Cape Flats, known mostly from collections in the 18th 90 M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100

Fig. 11. Known distribution of P. congesta: section of map shown is the south-western tip of South Africa.

and 19th centuries. Thus, based on the IUCN red list criteria (IUCN, Simons Bay (−AB), Wright s.n. (US); alongside Louwsrivier, Simons- 2012a,b)weclassifyP. aphylla as VU B2b(i,ii). town (−AB), Nov 1981, Stirton 9973, 9993 (K, PRE). Without precise locality: Anonymous 35 (P 04180816); Armstrong s.n. 4.2.5.5. Additional specimens examined. 3318 (Cape Town): Cape Town, (K, P); Peters s.n. (P); Verreaux s.n. (TCD); Zeyher s.n. (P 03496109); (−AB), Feb 1924, Smuts 1180 (PRE); Balston s.n. (BM); western Caput Bonae Spei, Burmann s.n. (G); Harvey 786 (BM); Harvey s.n. (K, streams of Table Mountain (−CD), 26 Oct 1924, Forbes 140 (PRE); MO); Jardin de Noisette, Gay s.n. (GB); Kataksy s.n. (OXF); 1840, Krauss Table Mountain (−CD), Deleaux 19 (P03496086); Deleaux s.n.(P, 902 (MO); Forster 1345 (BOL, K, PRE); Lehmann s.n. (P 03496088); Le 03496126); Ecklon & Zeyher 3545 (BOL); Ecklon 1828 (P); Lynes s.n. Vaillant s.n. (P 03496115); Miers s.n. (BM); Oldenberg s.n. (BM); Olden- (BM); 18 Nov 1897, Galpin 3965 (PRE); Sonnerat s.n. (P03496100); burg 104 (BM); Perrotet 12 (P); Sieber s.n. (L); Wallich s.n. (BM); Wallich Table Mountain, Nursery Valley (−CD), Martinez-Azorin 81 (NBG); s.n. (BM, K); Cape Flats, Rehmann 2015 (BM); Cape Peninsula, Salter between Cape Town and Table Mountain, on the plain (−CD), 5 Dec 7977b (BOL); Tafelberg C.B.S., Peters s.n. (BM, K, US, Z); Cape District, 1810, Burchell 49 (K); Bolus s.n. (TCD); Nov, Ecklon & Zeyher 1530 Bowie 8 & 9 (BM); Niven 159 (BM); Cape of Good Hope, Miers 165 (TCD); Harvey 251 (BM, MO, TCD); west side of Devils Peak (−CD), Al- (BM); Incorrectly labelled as Niligiri Hills India, Hohenacker 1198 (P). exander Prior s.n. (K); Nov, Ecklon & Zeyher 1519 (LE); Table Mountain (−CD), Nov 1892, Bolus 7298 (BOL, PRE); Sep, Ecklon 47 (K, OXF, PRA, 4.2.6. Psoralea congesta C.H.Stirt. & Muasya sp. nov. PRE), Oct, Ecklon 660 (PRE); Pappe s.n. (K); Lynes s.n. (BM); 20 Sep Psoralea congesta C.H.Stirt. & Muasya sp. nov., differs from P. aphylla 1958, Werderman & Oberdieck 42 (PRE); Table Mountain, Saddleback, in its stiff, grey stems, bearing many short compact broom-like short Devil's Peak (−CD), Oct 1886, Thode 8347 (PRE); Nov, Ecklon & Zeyher seasonal shoots aggregated in bunches at the apex, black-haired pedi- 1531 (BOL, SAM, TCD); Thode s.n. (PRE); Vlakkenberg, 26 Dec 1896, cels and calyces, congested appressed scales, with very short internodes, Wolley-Dod 2257, 2607 (BOL); Kirstenbosch (−CD), Van Wyk 2740 purple and white flowers and without vein colouring on the standard. In (JRAU). contrast, P. aphylla is a willowy shrub to small tree, with a few lax sea- 3418 (Simonstown): Kogelbaai (−AB), 27 Nov 1981, Stirton 9971 sonal shoots, silvery hairy pedicels and calyces, uncongested free scales, (K, NU); Chapman's Peak drive (−AB), 19 Oct 1986, Stirton 11212 (K, with long internodes, bluish/mauve flowers with dark purple veins; NBG); Rondeheuwel (−AB), 19 Oct 1986, Stirton 11206 (NBG); road Psoralea sp. 3, Stirton & Schutte in Manning & Goldblatt, Strelitzia 29: from Camps Bay to Hout Bay (−AB), 19 Dec 1927, Young 230 (PRE); 572 (2012). Type: South Africa, Western Cape, Simonstown (3418): Camps Bay (−AB), 10 Oct 1956, Cassidy 44 (BOL, NBG); kloof above Silvermine Nature Reserve, Crassula Ridge, Steenberg Plateau (−BC), Camps Bay (−AB), 7 Dec 2009, Muasya, Chimphango & Stirton 3400 5 Oct 2008, Dludlu & Stirton 39 (BOL!, holo.; NBG!, iso.). (BOL); 20 Sep 2009, Muasya & Stirton 4561 (BOL); 29 Jan 1908, Rogers Stiff shrubs to 2 m tall, reseeder, colonial. Stems 1–3, greyish black, 3022 (GRA); Ecklon 36 (OXF, PRA, PRE); Hoepoe Avenue, Camps Bay mostly arising from just above the base, rough; branches erect and (−AB), 8 Mar 2008, Muasya & Stirton 3892 (BOL); Cape Flats (−AB), spreading, stiff even when in flower. Seasonal shoots erect, densely ca- Ecklon s.n (PRE); near Sirkelsvlei (−AB), 6 Dec 1938, Salter 7894 nescent, becoming glabrous with age, broom-like, up to 40 cm long, (BOL); Modderdam, Cape Peninsula (−AB), 3 Dec 1938, Salter 7877b “burst branching” from last season's branch ends; leaves absent in ma- (BOL); Bergvliet Farm, along sluit at Firgrove Hill (−AB), Purcell s.n. ture plants, seasonal shoots covered in scales. Scales 2–3mmlong, (SAM 89905, 89906); Bergvliet Farm, near lake (−AB), Purcell s.n. 1.0–1.5 mm wide, green, persistent, becoming woody, subulate, tightly (SAM 89904); east of Plumstead (−AB), Salter 25618 (BM); Forbes 23 clasping and overlapping, densely white pubescent below but glabrous (BM); Cape Flats, Muizenberg (−AB), Lynes s.n. (BM); Wolley-Dod above, margins ciliate, densely minutely black gland-dotted when dry. 2334 (K); Sweet Valley (−AC), Wallich 94 (BM); H. Evans Santa Monica Inflorescences borne in uppermost 3–4 axils of seasonal shoots, con- Place (−AB), 10 Jun 1929, Walther s.n. (AA); Boyes Kraal River (−AB), densed, opening together, pseudo-capitate; buds hairy; pedicels below Compton 16662 (NBG); behind Simonstown (−AB), May 1923, Smuts cupulum 2–3 mm long, stiff and erect, hairy; bracts 2 mm cucullate; 725 (PRE); Simonstown Hill (−AB), Dec 1950, Pillans 10304 (MO); pedicels above cupulum 2–3 mm long; cupulum shallowly bilobed, M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100 91

ab c

d f

j o p

l m q k

Fig. 12. Vegetative and reproductive morphology of P. pullata: (a) cupulum opened up; (b) flower bud; (c) fruit; (d) cupulum; (e) mature flower; (f) scale; (g) standard petal; (h) standard petal folded; (i) wing petal; (j) keel petal; (k) calyx opened up: upper lobes to the left; (l) seed; (m) pistil; (n) stigma; (o) general habit; (p) seasonal shoot in flower; (q) close up of seasonal shoot. Voucher: Esterhuysen 2929 (BOL). Scale bars: a–m, 1 mm; o, 20 cm; p, 10 mm; q, 0.3 mm. Artist: A. Beaumont. overlapping the base of the calyx. Flowers 14–15 mm long, purple and vertical curved 2 mm long narrow white appendages present 2 mm mauve, veins not discoloured, maturing more or less simultaneously; above claw; nectar guide a M-shaped white area without purple flash; bracts inconspicuous, replaced by a tuft or ring of hairs. Calyx lobes auricles poorly-developed; apex emarginate. Wing petals shorter than calyx tube; lobes subequal, with lower lobe 1 mm longer; 11–13 × 4–5 mm, longer than keel, mauve, narrowly cultrate, lower lobe 4–6 mm long, 2.0–2.5 mm wide, cucullate, acute, broader upcurving; claw 4 mm long. Keel 9–13 × 3 mm, apex rounded; claw than other four equal lobes; vexillar calyx lobes free above the tube; 6 mm long. Androecium 11 mm long. Pistil 11 mm long, ovary 1.5 mm tube 3.0–4.0 mm long, covered in long appressed black hairs; calyx long; style filiform, 7 mm to point of flexure, glabrous but with a few shorter than corolla; inner face of calyx lobes densely covered in black stalked club-shaped glands on upper margin, pedicellate, length of cur- stubby hairs; calyx glands constant in size, same density all over, drying vature 4.5 mm, penicillate. Fruits 4.0–6.0 × 2.0–2.7 mm, papery, strongly orange; accrescent in fruit. Standard petal 12–13 × 10–12 mm, sub- reticulate, black, narrowly oblong. Seeds 2.0–4.0 × 1.3–2.0 mm, chestnut orbicular, purple fading to mauve at margins, veins not coloured but brown to black; hilum sub-central, aril well-developed, white, 1 mm drying black; claw 3–4 mm long, elongated and narrow, two free long (Fig. 10). 92 M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100

4.2.6.1. Diagnostic characters. Psoralea congesta resembles P. aphylla in dense branching at each node. Seasonal shoots erect; growing tips having clasping and persistent scales in the seasonal shoots, a bifid densely covered in black and white hairs, becoming glabrous, striate, cupulum, with both lobes non-bilabiate, which is situated just below with randomly scattered glands. Scales 3.5–4.0 × 3.0 mm, broadly or overlapping with the calyx tube. It differs from P. aphylla in its mostly obliquely ovate, clasping the shoots, glabrous, not tightly congested. In- leafless erect stiff grey stems bearing many short compact broom-like florescences borne in upper axils of seasonal shoots but without leafy ex- short seasonal shoots aggregated at the apex, black-haired pedicels tension; pseudo-spicate; lax; pedicel below cupulum 4–5mmlong, and calyces, congested appressed scales, short internodes; and purple stout and rigid; pedicel above cupulum 5–6 mm long; cupulum bilobed and white flowers without vein colouring. with one of the lobes variously bilabiate; lobes clasping and overlapping with calyx tube, equally developed and triangular, completely covered 4.2.6.2. Distribution and habitat. Psoralea congesta is distributed in the in long black hairs; glands of cupulum may be inconspicuous due to higher reaches of the Cape Peninsula and a few localities across False the dense pubescence. Flowers 15–17 mm long, mauve, maturing se- Bay (Fig. 11), at altitudes (500) 700–1000 m above sea level. This spe- quentially; bracts reduced to tuft or ring of hairs. Calyx tube 4 mm cies occurs mostly on western and eastern aspects of Mountain fynbos: long, completely covered in long black hairs; lobes equal, but longer Peninsula Sandstone Fynbos i.e., FFs 9 (Rebelo et al., 2006). Flowering than calyx tube; lateral and vexillar calyx lobes acute, straight, and tri- time is August to December. angular; lower lobe 5–7×3–5 mm, acute, as wide as other four lobes; vexillar calyx lobes fused for up to one third of their length above the 4.2.6.3. Etymology. The specificepithetcongesta is derived from the tube; calyx shorter than corolla; inner face of calyx lobes densely cov- Latin word congestus –a –um meaning ‘heaped together’ and refers to ered in long black hairs; ribs distinctly thickened; glands small, dense, the congested nature of the short seasonal shoots and congested scales. orange, constant in size and equally distributed across the calyx lobes and tube. Standard petal 14–15 × 11–15 mm, sub-orbicular; claw 4.2.6.4. Conservation status. This species is very common where it occurs 4–5 mm long, elongated and narrow; mauve; apex emarginate. Wing in small colonies and is less threatened than P. aphylla on the Cape petals 13.5–15.0 × 6.0 mm, lightly attached to keel, up-curving, longer Peninsula and it falls mostly within protected areas. Hence, based on than keel; claw 6 mm long; auricles swollen. Keel 11–13 × 4 mm; the IUCN red list criteria (IUCN, 2012a,b) we classify P. congesta as claw 7–8mmlong.Androecium 11 mm long, upper end curved inwards, Least Concern (LC). 10th stamen lightly attached, fenestrate. Pistil 11 mm long, ovary 2 mm long; style 7 mm long, swollen beyond point of flexure, length of curva- 4.2.6.5. Additional specimens examined. 3318 (Cape Town): Klawer Vlei ture 5 mm. Fruits 5 × 3 mm, membranous, reticulate. Seeds (−AB), 16 Jan 1896, Wolley-Dod 298 (BM, BOL, NBG); top of 4.0 × 2.5 mm, brown; hilum sub-central, aril well-developed, white Table Mountain (−BD), Oct 1887, MacOwan 924 (BM, BOL, GRA, K, (Fig. 12). SAM, US, Z); Drège s.n. (P 03496099); Ecklon s.n. (P 03050240); Garside 251 (K); Smuts track to left of Window Buttress (−BD), 31 Dec 2007, 4.2.7.1. Diagnostic characters. Psoralea pullata can be confused with Muasya & Stirton 3551 (BOL); slope over Wynberg ranges (−BD), 20 P. aphylla and P. congesta because all three species have their leaves re- Oct 1895, Wolley-Dod 21 (BOL); Skeleton Gorge (−CD), 13 Sep 2009, duced to scales. However, it can be distinguished by a combination of its Muasya & Stirton 4536 (BOL, NBG). spreading habit with pendulous branches, white hoary flowering shoot 3418 (Simonstown): Wynberg flats (−AB), Harvey 94 (TCD); tips, sweet-smelling mauve flowers, white wing petals, densely black- between Wynberg and Kalk Bay (−AB), 1873, Bolus 2607 (BOL); haired calyces and cupula, calyx lobes that are much longer than the Noordhoek Peak (−AB), 1 Nov 1942, Barker 2083 (NBG); Sandvlei calyx tube, and glabrous fruiting calyces. Psoralea congesta differs in its (−AB), Martin 4312 (GRA); Vergelegen Estate (−BB), 15 Oct 2011, stiff erect, rough, grey, multiple stems bearing many short compact Stirton 13378 (BOL, NBG); Disa Ridge, Silvermine (−AC), 28 Nov 1981, broom-like short seasonal shoots aggregated at the apex; congested, ap- Stirton 9974 (K, MO); Crassula Ridge, Silvermine Nature Reserve pressed and mostly overlapping scales, and in its deep purple flowers. (−BC), 10 Dec 2009, Muasya & Stirton 3408 (BOL); 10 Dec 2007, Stirton Psoralea aphylla has a similar habit but differs in its long bifid cupulum 13791 (BOL). that overlaps with the calyx, the tips of seasonal shoots which are 3419 (Caledon): Hottentots Holland, Caledon (−AB), 9 Sep 1928, covered with white silvery hairs, weakly scented purple flowers with Hutchinson 313 (BOL, K); Hermanus (−AC), 4 Oct 1924, Levyns 2591 darker veins, and white wing petals. (BOL); near Kleinmond turn-off to Caledon (−AC), 18 Sep 1949, Barker 5881 (BOL, NBG, STE); flats at Rooi Els (−AC), 20 Apr 1950, Parker 4470 4.2.7.2. Distribution and habitat. Psoralea pullata is a rare species that is (BOL). distributed from the Kogelberg Biosphere Reserve westwards to Without precise locality:Cap.Bon.Spei,Anonymous s.n. (P Potberg Mountain, with its main distribution in the Kleinrivier Moun- 06834348); Barbier s.n. (P 03496124); Boivan s.n. (P 03496102); Bunge tains (Fig. 13), at altitude 160–230 m above sea level. It occurs in seep- s.n. (P 03496095); Sieber 36 (P 03050210); Viellard s.n. (P 03496090); ages in Overberg Sandstone Fynbos i.e. FFs 12 (Rebelo et al., 2006). Viellard & Deplanche s.n. (P 03050209); Wanman 95 (SBT); near Smith's Flowering time is August to September, fruiting until December. farm, Salter 6561 (SAM). 4.2.7.3. Etymology. The name pullata is derived from the Latin pullatus, 4.2.7. Psoralea pullata C.H.Stirt sp. nov. meaning ‘clothed in black’ and refers to the distinctive black densely Psoralea pullata C.H.Stirt sp. nov., differs from P. aphylla and haired calyces. P. congesta in its multi-stemmed, floppy habit, larger flowers, densely black-haired calyces, and longer flower pedicels. Psoralea aphylla and 4.2.7.4. Conservation status. This species is abundant and widely distrib- P. congesta aresinglestemmed,stiff,smallflowered, with white and sil- uted in its habitat, but may be vulnerable due to habitat transformation ver haired calyces and short flower pedicels; Stirton & Schutte in on lower slopes for housing and agriculture. Therefore, based on the Goldblatt & Manning, Strelitzia 9: 506 (2000); Psoralea sp. 8, Stirton & IUCN red list criteria (IUCN, 2012a,b) we consider it to fall under the Schutte in Manning & Goldblatt, Strelitzia 29: 573 (2012). Type: South LC category for the time being. Africa, Western Cape, Caledon (3419): Vogelgat Nature Reserve, Fault Fall (−AD), 14 Feb 2010, Muasya & Stirton 5413 (BOL!, holo.). 4.2.7.5. Additional specimens examined. 3418 (Simonstown): Betty's Bay Large shrubs, 2–3 m tall, reseeder, occasionally forming dense (−BC), 5 Nov 1981, Snijman 569 (NBG); Kogelberg State Forest (−BD), clumps. Stems 1–3, woody throughout, weakly furrowed but generally 15 Sep 1992, Kruger 772 (NBG); Rooi Els (−BD), 2 Oct 1945, Leipoldt rounded; branches rigid, emerging in upper portions of plant, with 4189 (BOL). M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100 93

Fig. 13. Known distribution of P. pullata (squares), P. peratica (triangles) and P. ramulosa (circles): section of map shown is the south-western tip of South Africa.

3419 (Caledon): Kleinrivierberg (−AB), Aug 1940, Esterhuysen 2929 one third of their length above the tube; calyx shorter than corolla; (BOL); Fernkloof Nature Reserve (−AD), Van Wyk 1365 (JRAU); upper inner face of calyx lobes densely covered in small black stubby hairs; Fernkloof Nature Reserve (−AD), 30 Sep 1981, Bean 690 (BOL); Schutte ribs distinctly thickened; glands dense, constant in size and equally 50 (JRAU); east of Rocklands Peak, Klein Rivier Mountains, (−AD), distributed across the calyx tube and lobes. Standard petal Stokoe s.n. (SAM 55804); Vogelgat Nature Reserve, Hermanus (−AD), 14–16 × 12–18 mm, ovate-oblong; claw 4–6 mm long, elongated and 11 Nov 2007, Muasya & Stirton 3178 (BOL); Van Wyk 3204 (JRAU); narrow; auricles well-developed, large and swollen; apex rounded. Vogelgat (−AD), 9 Sep 1978, Williams 2566 (NBG, PRE); Fault Fall, Wing petals 14–18 × 5–7 mm, up-curving; claw 4–6 mm long; fused Vogelgat Nature Reserve (−AD), 14 Feb 2010, Muasya & Stirton 5413 to, but longer than keel. Keel 12–15 × 3–4 mm; claw 6–7mmlong.An- (BOL); Vogelgat Nature Reserve (−AD), intersection of Pearly Beach droecium 11–12 mm long, upper end curved inwards, 10th stamen road with Baardskeerdersbos/Elim road, Patterson-Jones s.n. (NBG); lightly attached; fenestrate. Pistil 11 mm long, ovary 2 mm long, sparsely 50 km from Elim/Bredasdorp road junction (−BA), 2 Nov 2008, Dludlu, glandular near apex, style 7 mm long, swollen at point of flexure, length Muasya & Stirton 71 (BOL). of curvature 4 mm, stipitate, penicillate. Fruits and seeds unknown 3420 (Bredasdorp): De Hoop, near Elandspad homestead (−BC), 16 (Fig. 14). Oct 1984, Van Wyk 2028 (NBG); Potberg (−BD), 21 Nov 1980, Stirton 8251 (PRE); west base of the Potberg (−BC), 13 Oct 1940, Pillans 9293 4.2.8.1. Diagnostic characters. This species is often confused with (BOL). P. aphylla, due to its lack of leaves. Psoralea peratica is characterised by Without precise locality: Zeyher s.n. (P 03496098). its pendant weeping habit with lax upper branches, densely velutinous branch tips and its large dark mauve flowers clustered at the ends of 4.2.8. Psoralea peratica C.H.Stirt short seasonal shoots. This very attractive species is restricted to the S. Afr. J. Bot. 64, 4: 244 (1998); Stirton & Schutte in Goldblatt & Piketberg Mountains. Helme 5225, collected on the SE slopes of Levant Manning, Strelitzia 9: 506 (2000); Stirton & Schutte in Manning & Hill, probably belongs here but differs in its glabrous calyces and stiffer Goldblatt, Strelitzia 29: 572 (2012). Type: South Africa, Western Cape, more erect habit. Meanwhile, P. aphylla is restricted to the Cape Penin- Clanwilliam (3218): western extremity of Piketberg, Gys-se-Kraal, sula, has erect upper branches and small, bluish-mauve flowers borne above Aurora (− DC), 19 Oct 1986, Rourke 1863 (NBG!, holo.; BOL!, in most axils of its seasonal shoots. K!, NBG!, PRE!, iso.). Large shrubs to small trees, 3–5 m tall, reseeder, may be colonial, but 4.2.8.2. Distribution and habitat. Psoralea peratica is a montane species, en- never forms dense clumps. Stems 1–3, erect, woody throughout, terete; demic to the Piketberg Mountains (Fig. 13) and occurs between 300 and branches flexuous, emerging at a height of about 1.0–1.5 m, arching. 1700 m above sea level. This species grows mainly along sandy seepages Seasonal shoots spreading or arching, densely pubescent; leaves absent and stream sides in Piketberg Sandstone Fynbos i.e. FFs 6 (Rebelo et al., in mature plants, seasonal shoots covered in scales. Scales 2.5–3.0 mm 2006). Flowering time is (September) October–November (February). long, persistent, clasping, glabrescent, not tightly congested. Inflores- cences borne in uppermost axils of bare seasonal shoots, pseudo- 4.2.8.3. Etymology. The epithet peratica comes from the Greek word capitate, congested, with 3–9 flowers per axil; pedicel below cupulum περάτίκός meaning “foreign, dwelling on the other side” and refers 2 mm long, stout and rigid; cupulum bilobed with one of the vexillar to the isolated distribution of this species in the genus. lobes variously bilabiate, hairy all over; cupulum lobes equally developed, glandular and narrowly triangular; pedicel above cupulum 4.2.8.4. Conservation status. The Red List of South African Plants (version 5–8mmlong.Flowers 16–18 mm long, dark mauve, maturing sequen- 2014.1; Raimondo et al., 2009) lists it as EN B1ab(ii,iii,iv,v). This species tially. Calyx tube 3 mm long, sparsely covered in white hairs and shorter has a narrow distribution range and is not very abundant where it oc- than the lobes; lobes more or less equal; lateral and vexillar calyx lobes curs. Helme and Raimondo (2008) report that “Subpopulations on acute, falcate and triangular; lower lobe 6–7 × 2.5–3.0 mm, acute, the lower slopes are declining due to ongoing habitat loss and degradation same width as the other four lobes; vexillar calyx lobes fused for up to due to crop cultivation, water extraction and overgrazing. 94 M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100

a b

h g j f

Fig. 14. Vegetative and reproductive morphology of P. peratica: (a) cupulum opened up; (b) cupulum; (c) seasonal shoot in ﬂower; (d) close up of seasonal shoot with scales; (e) calyx opened up: upper lobes to the left; (f) pistil; (g) standard petal; (h) keel petal (i) mature ﬂower; (j) wing petal. Voucher: Rourke 1863 (NBG). Scale bars: a, b, e–j, 1 mm; c, 10 mm; d, 3 mm. Artist: E. Ward-Hilhorst.

Subpopulations on upper slopes are relatively safe, apart from alien in- Mountains (−DA), 2 Dec 1981, Stirton 10058 (PRE); Platkloofrivier vasive plants becoming dense in some parts of the habitat and a long- Mountain (−DC), 23 Sep 1981, Stirton 9324 (PRE); Edwards 248 (BOL, term reduction in water supply as a result of a significant drying up of PRE); plateau east of Levant, Zebrakop (−DC), 20 Feb 1974, Linder 215 this area (rainfall has dropped 50% in 40 years according to farmers)”. (BOL); Gys-se-Kraal, above Aurora, western extremity of Piketberg (−DC), 19 Oct 1986, Rourke 1863 (NBG); south east slopes of Levant 4.2.8.5. Additional specimens examined. 3218 (Clanwilliam): New Cale- Hill, Piketberg (−DA), 1 Jan 2008, Helme 5225 (NBG). donia Farm, 20.3 km from R399 into Piketberg Mountains (−AD), 2 Nov 2008, Dludlu, Muasya & Stirton 72, 74 (BOL); Piketberg Mountains 4.2.9. Psoralea ramulosa C.H.Stirt sp. nov. (−AD), 16 Nov 2008, Dludlu, Muasya & Stirton 80 (BOL); Avontuur Psoralea ramulosa C.H.Stirt sp. nov., differs from P. usitata in its (−DA), 15 Dec 1981, Linder 3142 (BOL); Kapteinskloof, Piketberg brownish yellow stems up to 1.5 m tall covered in white lenticels; M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100 95 seasonal shoots smooth; leaf scales 1.2–2.5 mm long, pale mauve; profuse at each node and non-uniform, giving the plant an indetermi- flowers 17–18 mm long, borne in upper axils of seasonal shoots. nate architecture. Seasonal shoots glabrescent, with small scattered Psoralea usitata has greenish-grey to pale stems, 2.5–3.0 m tall, not prostrate hairs, spreading or arching, yellowish green, smooth, copious- lenticelled; seasonal shoots smooth, but with minute striations; leaf ly gland-dotted; glands drying black; leaves absent on mature plants scales 2.5–3.5 mm long, and flowers 9–14 mm long, borne in most but the basal nodes of watershoots and seedlings bear 1–2(3),lanceo- axils of seasonal shoots; Stirton & Schutte in Goldblatt & Manning, late, bright green, flat leaflets. Scales 1.2–2.5 × 0.8–1.0 mm, lanceolate, Strelitzia 9: 506 (2000); Psoralea sp. 9, Stirton & Schutte in Manning & clasping, glabrous, not tightly congested, persistent. Inflorescences Goldblatt, Strelitzia 29: 573 (2012). Type: South Africa, Western Cape, borne in upper axils of seasonal shoots but without leafy extension, Wuppertal (3219): Cederberg, 22 km from Clanwilliam towards pseudo-spicate, lax; pedicel below cupulum 3 mm long, stout and Algeria (−CA), 6 Dec 1981, Stirton 10200 (BOL!, holo.; NBG!, PRE!, iso.). rigid; cupulum bi-lobed with one of the vexillar lobes variously bilabi- Small shrubs, 1.0–1.5 m tall, resprouter. Stems two to three, ate, glabrous, lobes equally developed, narrowly triangular, glands con- branching densely from near the base, becoming woody throughout, spicuous; pedicel above cupulum 2–3mmlong.Flowers 12–15 mm brownish yellow with scattered white lenticels, rough surfaced, terete, long, pale mauve, maturing more or less simultaneously; bracts absent. not furrowed; branches flexuous, with randomly scattered glands, Calyx 7mmlong;tube3–5 mm long, glabrous, pale green, shorter than

g b h a

k e

Fig. 15. Vegetative and reproductive morphology of P. ramulosa:(a)matureflower (side-view); (b) flower bud; (c) scale; (d) cupulum; (e) calyx opened up: upper lobes to the right; (f) seasonal shoot in flower; (g) standard petal; (h) pistil; (i) androecium; (j) keel petal; (k) wing petal. Voucher: Stirton 10200 (BOL). Scale bars: a–e, g–k, 1 mm; f, 4 cm. Artist: A. Bello. 96 M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100 the lobes; lobes subequal, lower lobe slightly longer; lateral and vexillar east of Citrusdal (−CA), 4 Apr 1964, Hardy 1732 (K, PRE, STE); Kanje calyx lobes acute, falcate, lanceolate; lower lobe 4–5×2–3 mm, acute, Farm opposite Geelberg (−CA), 8 Feb 2009, Muasya & Stirton 4393 broader than other four lobes; vexillar calyx lobes free above the tube; (BOL); Middelburg (−CA), 8 Feb 2009, Muasya & Stirton 4395 (BOL); calyx shorter than corolla; inner face of calyx lobes finely covered in Jantjiesfontein, Citrusdal (−CA), 24 Nov 1995, Hanekom 2859 (BM, K, white stubby hairs; ribs slender; glands dense, constant in size, denser NBG); 2 km north of Citrusdal near Old Blacksmiths forge (−CA), 3 on the tube, often flushed purple. Standard petal 9–12 × 10–13 mm, Jun 1939, Salter 8124 (BOL). sub-orbicular; claw 4.0–5.5 mm long, elongated and narrow; mauve with prominent white appendages above the claw; apex emarginate. 4.2.10. Psoralea usitata C.H.Stirt sp. nov. Wing petals 12–16 × 3.0–5.5 mm; mauve to white, strongly up- Psoralea usitata C.H.Stirt sp. nov., differs from P. aphylla by its curving; claw 4.0–5.5 mm long; fused to and longer than keel. Keel resprouting habit (rarely reseeding); glabrous shoots; leaves reduced 11–13 × 3–4 mm; claw 5–7mmlong.Androecium 11 mm long, broadly to appressed, bright green, glabrous scales, trifid cupulum and glabrous fenestrate, upper part recurved. Pistil 10 mm long, ovary 3 mm long; calyx tubes. Meanwhile, P. aphylla is a reseeder, seasonal shoots densely style filiform, 6 mm to point of flexure, swollen at point of flexure, covered in silvery hairs, leaves reduced to free, green silvery hairy length of curvature 4 mm, glabrous but with many small stalked club- scales; cupulum bifid; calyx tube covered in black and white hairs; shaped glands on upper part, pedicellate, penicillate. Fruits and seeds Psoralea sp. 12, Stirton & Schutte in Manning & Goldblatt, Strelitzia 29: unknown (Fig. 15). 573 (2012). Type: South Africa, Western Cape, Cape Town (3318): Groot Drakenstein (−DD), 3 Mar 1951, Gray s.n. (BOL!, holo.). 4.2.9.1. Diagnostic characters. Psoralea ramulosa is endemic to the Large shrubs, 2.5–3.0 m tall, resprouter or reseeder, forming dense Cederberg Mountains. It may be confused with the variable species clumps in seepages. Stems solitary to many, greenish grey to pale tan, P. usitata (both species have glabrous seasonal shoots and calyces), woody throughout, terete, shallowly furrowed; branches arching, which in its distribution range differs in its 2–3 m tall greenish grey emerging in middle portions of plants. Seasonal shoots erect, smooth, stems; seasonal growth comprised of multiple shoots arising from glandular, with minute striations, bearing up to seven short side each node giving the plant a scruffy appearance; 2.5–3.5 mm long leaf branches; leaves usually absent in mature plants, if present, trifoliolate, scales; and 9–11 mm long purple and white flowers borne in most 13–17 (20) × 4 mm, flat, covered in minute glands, lateral leaflets small- axils of seasonal shoots. Although P. ramulosa may resemble P. rigidula er. Scales 2.5–3.5 × 0.5–1.7 mm, senescent, appressed but spreading on herbarium sheets due to its glabrous seasonal shoots and calyces, when dry, not tightly congested. Inflorescences axillary, pseudo- its growth form and flower colour are very distinct. Psoralea rigidula spicate, lax; pedicel below cupulum 2–4 (5) mm long, stout and rigid; has numerous stiff herbaceous stems, never grows taller than 0.6 m cupulum bilobed with one of the vexillar lobes strongly bilabiate, cleft and has purple and white flowers, whereas P. ramulosa has pale to half the length of vexillar lobes and appearing trifid; cupulum gla- mauve flowers and two to three 1.0–1.5 m high stems that branch pro- brous, lobes equally developed, narrowly triangular; pedicel above fusely from near the base. The two species are also allopatric. cupulum 2–3 mm long, hairy. Flowers 9–14 mm long; purple and white; maturing sequentially; bracts reduced to tuft or ring of hairs. 4.2.9.2. Distribution and habitat. Endemic to the Cederberg Mountains Calyx 5–6 mm long; tube 2.5–3.0 mm long, glabrescent, longer than (Fig. 13), where it occurs mostly in scattered colonies and the dark the lobes; lobes subequal, the lower lobe slightly longer; lateral and green bushy plants look very distinctive scattered across hillsides. vexillar calyx lobes acute, falcate, narrowly triangular; lower lobe Psoralea ramulosa is restricted to Cederberg Sandstone Fynbos i.e. FFs 2.5–3.0 mm long, 1.5–2.5 mm wide, acute, broader than the other four 4(Rebelo et al., 2006)at155–1090 m above sea level. It may be found lobes; vexillar calyx lobes fused for up to one third of their length above along stream banks and in rocky seepages, but it occurs mostly on the tube; calyx shorter than corolla; ribs distinctly thickened; glands drier hillsides. Flowering time is November to April with a peak from dense, drying reddish, constant in size and equally distributed across December to February. The flowers are visited by Megachilid bees. the lobes and tube. Standard petal 10–12 × 11–13 mm, sub-orbicular; claw 2–5 mm long, elongated and narrow; apex emarginate. Wing petals 4.2.9.3. Etymology. The specific epithet ramulosa derives from the Latin 11–16 × 4–6 mm, up-curving; auricles present; claw 3.5–5.0 mm long; word ramulosus (many small, veining branches or twigs) and refers to fused to, but longer than keel. Keel 12 × 3–4 mm; claw 4–6mmlong.An- its dense habit. droecium 11 mm long, upper end curved inwards, 10th stamen lightly attached, fenestrate. Pistil 9 mm long, ovary 2 mm long, style 7 mm long, 4.2.9.4. Conservation status. Psoralea ramulosa has a narrow distribution swollen at point of flexure, length of curvature 3 mm. Fruits 3×3mm,pa- range and a small area of occupancy (b2000 km2), hence based on the pery, reticulate. Seeds 3mmlong,2.5–2.5 mm wide, brown; hilum sub- IUCN criteria (IUCN, 2012a,b) its present status corresponds to VU terminal, aril poorly-developed, white (Fig. 16). B2b(i,ii). 4.2.10.1. Diagnostic features. Psoralea usitata, like all other members of 4.2.9.5. Additional specimens examined. 3218 (Clanwilliam): 22.5 km the P. aphylla complex is often identified as P. aphylla due to its lack of from Citrusdal to Clanwilliam (−AC), 12 Dec 1981, Grobbelaar 2675 leaves in mature plants. However, it differs from the true P. aphylla by (PRE); Rietvlei Farm near Keurbos between Algeria and Clanwilliam a combination of sparsely hairy to glabrescent shoots and glabrous on riverbanks (−BD), 7 Feb 2009, Muasya & Stirton 4357 (BOL); Jan calyx tubes, sparsely hairy pedicels, small flowers, bifid cupulum with Dissels Rivier up to source (−CC), 18 Jan 1947, Leipoldt 4385 (K, PRE). vexillar lobe cleft and appearing trifid, and leaves reduced to appressed, 3219 (Wuppertal): Uitkyk Pass, Cederberg (−AA), 7 Sep 2008, bright green, glabrous scales. Muasya & Stirton 3944 (BOL); Cedarberg (−AC), 7 Aug 2009, Dludlu, Psoralea usitata is a variable species and there are four distinct vari- Muasya & Stirton 24 (BOL); top of Uitkyk Pass (−AA), 7 Feb 2009, ants. The typical form is an erect reseeding plant similar in habit to Muasya & Stirton 4363, 4368, 4369 (BOL); near Algeria Forest Station P. aphylla but with greenish brown stems and pale mauve flowers. The (−AC), Stokoe s.n. (SAM 55800); 1 km from Uitkyk Pass on road from standard petals have mauve limbs with purple veins and in the centre, Algeria to Cederberg (−AC), 7 Feb 2009, Muasya & Stirton 4362 (BOL, at the base at the point of flexure there is a white nectar patch under a K, NBG, PRE); Dasklip Pass, Beaverlac Nature Reserve (−AC), 12 Dec dark purple flash (Form 1: Type). Form 2 is a resprouter restricted to 2007, Muasya & Stirton 3414 (BOL); Middelberg Pass, Middelberge the Cederberg Mountains and is characterised by its tall untidy floppy (−AC), 8 Feb 2009, Muasya & Stirton 4395 (BOL); Donkershoek Kloof, habit with seasonal burst-branching from stem tips and along the old Cederberg (−CA), Stokoe s.n. (SAM 52303); east of Citrusdal, near greyish wood. The rarer Form 3 (Hanekom 1776, Helme s.n.) occurs in farm Allandale (−CA), 13 Nov 1985, Snijman 973 (NBG, PRE); 6 km the mountainous eastern parts of the Cederberg and is a stiff erect M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100 97

b a h

c i

d j

f l

Fig. 16. Vegetative and reproductive morphology of P. usitata:(a)matureflower (side-view); (b) scale; (c) mature flower (front-view); (d) cupulum; (e) flower bud; (f) calyx opened up: upper lobes to the right; (g) seasonal shoot; (h) standard petal; (i) pistil; (j) androecium; (k) keel petal; (l) wing petal. Voucher: Gray s.n. (BOL). Scale bars: a–f, h–l, 1 mm; g, 5 cm. Artist: A. Bello.

resprouting 5–6-stemmed plant with tall bare stems and densely clus- habit. Form 4 is characterised by its heavily lenticelled yellowish tan tered seasonal shoots at the apex (like witches brooms). The deep pur- stems, mauve to blue standard petals with a central purple flash, ple standard petals contrast strongly with the large white wing petals. white wing petals, and its distinctive “pom-pom” mass-flowering inflo- Form 4 (Psoralea sp. 5; Stirton and Schutte, 2012) is a lowland reseeding rescences during the flowering season. The plants produce numerous plant that is common throughout the Hex and Breede River valleys. The seasonal shoots; each of which terminates in 5–6 flowers. There are young plants look very similar to P. aphylla and Form 1 during the first often so many flowers produced that the plants become over-laden few years of growth but as they age they develop a very different and the branches droop resembling “pom-poms”.Furtherfieldwork 98 M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100

Fig. 17. Known distribution of P. usitata: section of map shown is the south-western tip of South Africa. may show that some of these forms may deserve infraspecific or specific 3319 (Worcester): Slopes of Drostersberg, upper catchment of Groot status. More collections are needed from the Cederberg region and Kliphuisrivier, Groot Wintershoek, 1.7 km from waterfall (−AA), Louw southern coastal areas from Gansbaai to De Hoop. 4418 (BOL); Groot Winterhoek – Groot Kliphuis River/24 rivers (−AA), Louw 4191 (BOL); Porterville Mountains (−AA), 16 Mar 1974, 4.2.10.2. Distribution and habitat. This species is widespread within the Thompson 2030 (NBG, PRE, STE); Witelsrivier (−AA), 16 Mar 1997, Western Cape Province (Fig. 17), particularly along mountain ranges. Low 3465 (BOL); trail to Die Hel, Groot Wintershoek Mts (−AA), 31 It does not overlap with P. aphylla but is sympatric with P. rigidula and Jun 2009, Muasya & Stirton 4340, 4343 (BOL, NBG, PRE), 4344, 4347, P. fleta in parts of their range. It occurs mostly along streams and seep- 4349 (BOL); Bainskloof (−AB), 14 Nov 1986, Schlechter 1246 (PRE); 5 ages in mountain and lowland fynbos in the following vegetation Oct 2008, Dludlu 36 (BOL); Schlechter 9183 (BM); 7 Feb 1966, Marsh types: FFb 2, FFg 2, FFs 5, FFs 10, FFs 11, FFs 12, FFs 14, and FFs 16 27 (K, NBG, STE); Garside 248 (K); Schelpe 1246 (PRE); 17 Jan 1945, (Rebelo et al., 2006), but also occurs on drier slopes, at altitude Compton 16917, 6 Apr 1946, Compton 17949 (NBG); 31 Nov 2007, 170–1600 m above sea level. Flowering occurs from September to May. Muasya & Stirton 3391 (BOL); Agterwitzenberg (−AC), 8 Mar 1966, Marsh 43 (K, NBG, PRE, STE); Sevenfontein Nature Reserve, Wolseley 4.2.10.3. Etymology. The specificepithetusitata is derived from the Latin (−AC), 26 Feb 1936, Hubbard 327 (NBG, STE); eastern slopes on top word usitatus –a –um (adj.) meaning common or ordinary and reflects of Mitchell's Pass, south of Ceres (−AC), 5 Apr 1974, Goldblatt 1346 its widespread occurrence. (MO, NBG, PRE); Tulbagh (−AC), Nov 1919, Rogers 17395 (PRE); Tulbagh Kloof (−AC), 14 Feb 1922, Pole-Evans 485 (PRE); Harvey s.n. 4.2.10.4. Conservation status. This species is very abundant and widely (BOL); Mostertshoek Waterfall, Ceres (−AD), 17 Jan 1944, Wasserfall distributed across a wide range of habitats. Therefore, based on the 829 (NBG); Haalsneeukop, Stokoe s.n. (SAM 56311); between Veepos IUCN catergories (IUCN, 2012a,b), we classify it as LC. and Groot Kliphuis, Great Wintershoek Mountains (−BB), 12 Dec 2007, Stirton & Muasya 3415 (BOL); south of Bainskloof Village, east of 4.2.10.5. Additional specimens examined. 3318 (Cape Town): above farm hut on Boland Hiking trail (−CA), 13 Dec 2007, Muasya & Stirton 3440 Waterval, western slopes of Kasteelberg (−BD), 15 Mar 2008, Helme (BOL); Mountains north of Bainskloof Pass (−CA), 22 Jun 2010, Muasya 5431 (NBG); Modderkloof, west side of Paardeberg (−DB), 14 Apr 5342 (BOL); Lewis 912 (SAM 56360); Slanghoek Peak (−CA), 2 Jan 1891, Hugo 2594 (PRE); Wellington (−DB), Nov 1882, Knobel s.n. 2001, Low 6837 (BOL); Elandspadrivier (−CA), 6 Jan 2003, Low 8326 (PRE); 4 May 1954, Van der Merwe 8 (NBG, STE); Sep, Cummings 21 (BOL); Elandskloof Valley SE of Voëlvlei Dam (−CA), 15 Jan 2008, Low (NBG); Berg Rivier, Klein Drakenstein Mountains, (−DB), Drége s.n. 15682 (BOL); Witterivier Valley, (−CA), 6 Apr 1959, Esterhuysen (TCD); Modderkloof, west of Wellington, 17 Jun 1981, Fellingham 21 28296 (BOL); Du Toits Kloof (−CA), Drège s.n. (BM, K); Stokoe s.n. (NBG); Jonkershoek Catchment area (−DC), 28 Jun 1946, Strey 620 (SAM, PRE); 22 Mar 1959, Esterhuysen 28237 (BOL, K); 10 Feb 2008, (PRE); 13 Feb 1949, Morris 3 (NBG); 17 Feb 1972, Van der Walt 180 Helme 5417 (NBG); 19 Jan 1975, Walters 1353 (NBG); farm (STE); 14 Feb 1966, Grobbelaar 393 (PRE); 28 Nov, 1960, van der Eensgevonden, Rawsonville (−CB), 24 Jul 2010, Muasya & Stirton Merwe 2119 (PRE); 14 Jan 1948, Rodin 3268 (BOL, K, MO, P, PRE); 27 5391 (BOL); Fairy Glen, lower slopes of Brandwagberge (−CB), 18 Sep Mar 1964, Bos 1386 (PRE); May 1967, Kerfoot 5787 (STE); 7 Jan 1976, 1981, Stirton 9142 (PRE); Waaihoek turnoff on Worcester–Ceres Road Kruger 144 (PRE); Maguire 3 (NBG); Nov 1962, Morris 46 (NBG); 24 (−CB), 3 Feb 1981, Walters 2426 (NBG); Worcester (−CB), 18 Oct Jun 2010, Muasya & Stirton 5364 (NBG); Jonkershoek Catchment area 1921, Fine 30 (PRE); Rehmann 2429 (NBG, STE); Groenrivier (−CB), 4 (−DD), 16 Nov 2007, Muasya & Stirton 3194 (BOL); Paarl Mountain Dec 1981, Stirton 10123 (NU); Rawsonville (−CC), 14 Dec 1938, Louw Nature Reserve (−DC), 29 Oct 2009, Muasya & Stirton 4846 (BOL); 105 (NBG); upper Zachariashoek Catchment (−CC), 20 Feb 1970, Waterkloof Farm (−DD), 29 Mar 1994, Nel & Boucher 75 (BM, K, Haynes 298 (NBG, PRE); Wemmershoek (−CC), Apr 1924, Smuts 1122 NBG); Verlorenkloof, Stellenbosch (−DD), Feb 1967, Geertsma 38 (PRE); Kasteelkloof, Worcester (−CC), 8 Mar 1973, Smith 101 (PRE, (STE); Groot Drakenstein (−DD), 3 Mar 1951, Gray s.n. (BOL); STE); Kliphuis (−CC), 9 Oct 2010, Muasya & Stirton 5690 (BOL); Swartboskloof (−DD), 28 Nov 1960, Van der Merwe 2119 (PRE). Theewaterskloof (−CC), 20 Feb 1929, Hugo 1489 (STE); Zuurvlakte M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100 99

189, northern Groot Winterhoek Mountains (−CC), 1 Mar 2008, Helme SE slopes of Kanonberg, Riviersonderend mountains above Greyton 5385 (NBG); 1.5 km S of turnoff to Mt. Rochelle Forestry area (−CC), 26 (−BB), 25 Jan 2008, Helme 5297 (BOL); top of the mountains of the Jan 1990, Steiner 2069 (NBG); Bainskloof (−CD), 22 Jun 2010, Muasya Baviaanskloof near Genadendal (−BB), 16 Feb 1815, Burchell 7738 5346 (BOL); 15 Dec 2008, Muasya & Stirton 4330 (BOL); Matroosberg (BM, K); Roser s.n. (TCD); Dashung near Stormsvlei (−BB), Oct 1947, MCA (−CD), Compton s.n. (PRE); Drakenstein Mountains (−CC), Stokoe Stokoe 55186 (PRE); 3 km N of Riviersonderend (−BB), 19 Nov 1980, s.n. (SAM); halfway down Franschoek Pass on northern side (−CD), Stirton 8219 (PRE); Vooruitzight, Riviersonderend Mountains (−BB), Stirton 13301 (BOL); Franschoek Pass (−DA), 12 Dec 1950, Compton 18 Apr 1995, Rourke 2077 (PRE); Paardekloof (−BC), 30 Mar 1970, 21904 (NBG); 22 Nov 1985, Goldblatt 7380 (MO, NBG); Kanetvlei, 26 Theron & Van der Schijff 2275 (PRE); Schoenmakerskloof Farm, between Oct 1983, Zeeman 17 (PRE); Great Wintershoek Reserve (−DB), 10 Kleinberg and Bredasdorpberge (−BD), 20 Nov 1980, Stirton 8220 (NU, Oct 2009, Muasya & Stirton 4641 (BOL); Modderkloof, west side of PRE); Faery Glen (−BD), 7 Dec 1960, Gentry 18792 (US); Zondereind Paardeberg, Paarl district (−DB), 14 Apr 1981, Hugo 2594 (STE); Paarl Mountains (−CC), Wilson 6 (GB). Mountain (−DB), Mar 1926 Grant 2209 (NBG, PRE); 17 Jan 1963, Kruger 3420 (Bredasdorp): National Bontebok Park (−AB), Nov 1962, 179 (PRE); mid N slopes of Jonaskop, between Worcester and Marais, 36 (PRE); Swellendam Forestry Road Reserve (−BA), 18 Feb Genadendal (−DD), Patterson-Jones 61 (NBG); Stellenbosch (−DD), 1971, Negin 16 (NBG); Swellendam Zuurbraak (−BA), 1 Mar 1930, 14 Mar 1957, Basson 16 (NBG); May 1967, Kerfoot 5787 (STE); Thode A2317 (PRE); Strawberry Hill, Heidelberg (−BB), Stokoe (PRE, Helshoogte (−DD), Apr 1927, Lotsy & Goddijn 1916 (L). SAM 61583, STE); Zuurbraak, Swellendam District (−DA), Jan 1930, 3320 (Montagu): Marloth Nature Reserve (−BD), 25 Oct 2008, Thode 2317 (K, PRE); Sandberg Farm, Elim region (−DA), 26 Nov Dludlu, Stirton, Chimphango & Muasya 40 (BOL); Tradouw Pass (−DC), 2011, Muasya & Stirton 13569 (BOL). Stirton & Žantovská 11638 (K, MEL,NU); Duiwenhoeksrivier at 3421 (Riversdale): mountains above Corente Rivier, Riversdale Zeekoeigat, Heidelberg Division (−DD), 20 Nov 1972, Bremer 541 District (−AA), Nov 1908, Muir 80 (PRE); Riversdale, second stream (BM, K); Goedehoop Farm close to banks of Duiwenhoeks River, flowing into Corente river (−AB), 14 May 1979, Bohnen 5692 (PRE); 6 Grootvadersbosch area (−DD), 27 Jun 1988, McDonald 1566 (NBG); Apr 1926, Smith 2775 (K, PRE); 26 Nov 1892, Schlechter 1881 (BM, Grootberg Mountain, Lemoenshoek Peak (−DD), 6 Dec 1941, Stirton GRA, PRA, PRE, TRVF1246). 10243 (K, NU); Between Lemoenshoek and Naauwkrantz Farm, Straw- Without precise locality : Drége s.n. (MO); Langley Kitching s.n.,South berry Hill, Langeberg (−DD), Stokoe s.n. (NBG, PRE); Lemoenshoek Africa (K); Caput Bonae Spei, Masson s.n. (BM); Niven 159 (BM); Peak (−DD), Stokoe s.n. (SAM, STE). Lamprechtsbos, Sep 1962, Taylor 3992 (PRE, STE); Houtech terrain, 3321 (Ladismith): Garcia's Pass (−CC), 8 Mar 2009, Muasya & Houwhoek, below Haasvlakte, 1 Dec 1987, Boucher 5267 (NBG). Stirton 4473, 1 Dec 2009, Muasya & Stirton 5087 (BOL); Garcia's Pass, near Toll House (−CC), 19 Apr 1983, Fellingham 443 (BM, K, PRE); 7 Acknowledgements Mar 1987, Bohnen 8770 (NBG); Muiskraal (−CC), Campbell 485 (NBG); Gysmanshoek Pass, south side (−CC), 25 Nov 1987, Rourke We acknowledge the South African National Biodiversity Institute 1900 (NBG); Wadrift Gate, Gysmanshoek Pass Road (−CC), 25 Nov (SANBI) — Threatened Species Programme (TSP) and the National 1987, McDonald 1488 (NBG, PRE); 16 Sep 1981, van Wyk 724 (PRE); Research Foundation (NRF-SABI — # 68793, AMM) for providing re- The Oaks farm, Gysmanshoek Pass (−CC), Van Wyk 74 (K, NBG); 10 search funding. MND thanks the following organisations for financial Dec 1981, Stirton 10284 (NU); Top of Cloete Pass, Herbertsdale support: the University of Cape Town; through its International Student (−DD), 15 Nov 1987, Vlok 1880 (NU, PRE, SAAS). Scholarship and the Dorothy Cameroon Scholarship, Canon Collins 3322 (Oudtshoorn): Moeras Rivier, 8.3 m from George (−CC), 27 Trust, and the University of Swaziland. CHS thanks Angus & Anne Nov 1943, Fourcade 6281 (BOL,NBG,PRE,STE). Bean (Claremont, Cape Town) for their valuable support during these 3418 (Simonstown): Sir Lowry's Pass (−BA), 20 Jan 1894, Kuntze studies. s.n. (BM, K); Stokoe s.n. (PRE); Hottentots Holland Mountains (−BB), We are grateful to a number of landowners, botanists and ecol- Stokoe s.n. (PRE 55184); Steenbras (−BB), Jan 1914, Rogers 10510 ogists for their interest and support: Nick Helme, Barrie Louw, Dave (BOL, K); Helderberg (−BB), 21 Sep 1928, Gillett 657 (STE); MacDonald, Gerhard van Deventer, Anso le Roux, Ishmail Ibrahim, Moordenaarskop (−BB), Stokoe s.n. (PRE); 13.7 km from Stanford to Rupert Koopman, Jan Vlok and Anne-Lise Schutte, Jane & Douglas Elim (−BC), 2 Feb 2011, Stirton & Muasya 13183 (BOL); Kogelberg Hill and Sally & Douglas Scott. We are also grateful to the curators State Forest (−BD), 24 Mar 1992, Kruger 446 (PRE); Platberg, head of and staff of the following 33 herbaria (BM, BOL, E, GRA, JRAU, K, L, Oudebos Forest (−BD), 20 Jun 1969, Boucher 385 (K, PRE, STE); near LE, LINN, M, MO, NBG, NH, NU, NY, OXF, P, PRA, PRC, PRE, PRU, S, weir, Han se Kop, Hottentots Holland Mountains (−BB), 2 Oct 2009, SAAS,SAM,STE,TCD,US)foraccesstoorloanoftheirmaterial. Muasya & Stirton 4753, 4755 (BOL). This study was undertaken under a number of different permits 3419 (Caledon): Farm Donderboskop (−AA), 4 Oct 1975, Walters from Cape Nature (AAA008-00035-0028 02.09.2011: C.H. Stirton; 1408 (NBG); Diepgat (−AA), 25 Nov 1981, Stirton 9949 (NU); Elgin 0028-AAA005-00161 31.12.2014: S. Chimphango); Department of (−AA), 6 Mar 1926, Smith 2565 (K, PRE); Lebanon Forest Reserve Environmental Affairs (K 135091); and the Provincial Administration (−AA), 7 Jan 1976, Kruger 144 (STE); 20 Dec 1966, Van der Zel 35 of the Cape of Good Hope (102/1984; 222/1985; 323/1985; 72/1986). (PRE); Viljoen Pass, Grabouw (−AA), 20 Dec 1956, Strey 2901 (PRE); We are grateful to the various environmental and conservation agencies Balfour-Browne 7 (BM); Lebanon river bed, Grabouw (−AA), 20 Dec for their permission to study material in situ on their properties. Finally, 1966, Van der Zel 235 (PRE); Kleinmond (−AA), 9 Nov 1981, Barker we would like to thank two anonymous reviewers who made some 1389 (NBG); Jakkalsrivier, Lebanon State Forest (−AA), 21 Nov 1972, vital comments and suggestions on an earlier version of the manuscript. Kruger 1600 (PRE); 8 Jan 1971, Kruger 1098 (PRE); Mt. Lebanon, Elgin (−AA), 7 Feb 1982, Esterhuysen 35746 (BOL); Caledon (−AA), Jan References 1968, Sidey 4113 (PRE); Hermanus (−AC), 25 Aug 1930, Sutton 447 (PRE); Stanford, road to Hermanus (−AC), 3 Dec 1986, Germishuizen Dludlu, M.N., Stirton, C.H., Chimphango, S.B.M., Bello, A., Muasya, A.M., 2013. Phylogenetic fl − position of the southern African members of the tribe Psoraleeae based on molecular 4120 (PRE); 5 Oct 1924, Levyns 2591 (BOL); Kleinmond ats ( AC), – − and morphological data. South African Journal of Botany 89, 150 155. 14 Sep 1946, De Vos 72 (STE, NBG); Mossel Rivier ( AC), Guthrie s.n. Egan, A.N., Crandall, K.A., 2008. Incorporating gaps as phylogenetic characters across eight (PRE); 12 km from turnoff into R320 from Hermanus to Caledon DNA regions: ramifications for North American Psoraleeae (Leguminosae). Molecular (−AD), 4 Apr 2004, Bester 5060 (PRE); Voëlklip (−AD), Williams 121 Phylogenetics and Evolution 46, 532–546. − Egan, A.N., Reveal, J.L., 2009. A new combination in Pediomelum and a new genus, (BM, K, NBG); Shangri-la, Maanskyn Reserve ( AD), 27 Jan 1988, Ladeania, from western North America (Fabaceae, Psoraleeae). Novon 19, 310–314. Williams 3808 (BM, K, NBG, NU); Vogelgat (−AD), Stokoe s.n. (PRE); Forbes, H.M.L., 1930. The genus Psoralea Linn. Bothalia 3, 1–35. 100 M.N. Dludlu et al. / South African Journal of Botany 97 (2015) 77–100

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