(Asparagaceae, Scilloideae), Including the Description of Desertia Luteovirens and the Taxonomic Revisions of Whiteheadia and Namophila

Home , Daubenya, Drimiopsis, Drimiopsis maculata, Eucomis, Ledebouria, Namophila

Phytotaxa 221 (3): 201–225 ISSN 1179-3155 (print edition) www.mapress.com/phytotaxa/ PHYTOTAXA Copyright © 2015 Magnolia Press Article ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.221.3.1

Desertia, a new genus in Massonieae (Asparagaceae, Scilloideae), including the description of Desertia luteovirens and the taxonomic revisions of Whiteheadia and Namophila

MARIO MARTÍNEZ-AZORÍN1,2,*, MICHAEL PINTER1 & WOLFGANG WETSCHNIG1 1Institute of Plant Sciences, NAWI Graz, Karl-Franzens-University Graz, Holteigasse 6, A-8010, Graz, Austria; e-mail: [email protected] 2dCARN & CIBIO (Instituto Universitario de la Biodiversidad), Universidad de Alicante, P.O. Box 99, E-03080 Alicante, Spain. *author for correspondence

Abstract

Desertia gen. nov., which belongs to tribe Massonieae, is described from south western Namibia and north western South Africa. Desertia is at first sight related to Whiteheadia and Massonia, but it can be clearly differentiated by the papillose bracts, subcampanulate flowers with straight, suberect or slightly spreading free portion of tepals and the rugose seeds. This genus is based on Whiteheadia etesionamibensis, for which the combination in Desertia is presented. Furthermore, a second species in the genus, D. luteovirens, is here described based on distinct morphological and molecular characters. Further- more, taxonomic revisions are presented for Namophila and Whiteheadia, including identification keys for the three genera and data on morphology, ecology, and distribution for all accepted species.

Key words: Flora of Southern Africa, Hyacinthaceae, Massonieae, Massonia, taxonomy

Introduction

Subfamily Scilloideae tribe Hyacintheae is alternatively treated as Hyacinthaceae subfam. Hyacinthoideae, a treatment that we favour here (Speta 1998a, 1998b, Wetschnig et al. 2002, Pfosser et al. 2003, Manning et al. 2004). Within subfamily Hyacinthoideae, the tribe Massonieae has shown important changes regarding generic circumscription in the last decades. Two new genera—Namophila Müller-Doblies & Müller-Doblies (1997: 77) and Spetaea Wetschnig & Pfosser (2003: 87)—were recently described whereas some others were included in broader generic concepts, such as Brachyscypha Baker (1870: 393), Polyxena Kunth (1843: 294), and Periboea Kunth (1843: 292) into Lachenalia J.Jacquin ex Murray (1784: 314) (Manning et al. 2004); Drimiopsis Lindley & Paxton (1851–1852: 73, fig.172) and Resnova Van der Merwe (1946: 46) into Ledebouria Roth (1821: 194) (Manning et al. 2004); Androsiphon Schlechter (1924: 147), Amphisiphon Barker (1936: 19) and Neobakeria Schlechter (1924: 149) into Daubenya Lindley (1835: t. 1813) (Manning & Van der Merwe 2002); and Whiteheadia Harvey (1868: 396) into Massonia Houttuyn (1780: 424) (Manning et al. 2004, 2011). The cited lumping of genera by Manning et al. (2004) was based on a preliminary phylogenetic study, including two plastidial regions (sequences not yet deposited in the Genbank) and an incomplete sampling of taxa in most of the groups studied. On this matter, Lebatha (2004) commented that “Manning et al. (2004) prematurely, without support of morphological data, sunk Drimiopsis and Resnova into Ledebouria sensu lato”. Furthermore, Lebatha (2004) added that her results “directly oppose Manning et al.’s (2004) decision to lump Resnova and Drimiopsis in Ledebouria based solely on a polytomy produced by a strict consensus tree of trnL-F and rbcL data. In addition, the characters professed by them to support their lumping prove to be either homoplasious, symplesiomorphous or confined to the terminal clade in Ledebouria”. Similar conclusions were reached by Lebatha et al. (2006) and by Martínez-Azorín et al. (2011) for Ornithogaloideae. Therefore it appears necessary to evaluate those proposals on the basis of extended phylogenies with additional regions and covering most of the species involved in each genus, and everything combined with a detailed analysis of morphological characters (M. Martínez-Azorín and collaborators, in preparation). Harvey (1868) described the genus Whiteheadia to include a single species, W. latifolia Harvey (1868: 396), based

Accepted by Lorenzo Peruzzi: 18 Jul. 2015; published: 4 Aug. 2015 201 on material collected by Henry Whitehead at Modderfontein, west of Springbok in northern Namaqualand, South Africa. However, Harvey (1868) was not aware that Jacquin (1791) had previously described Eucomis bifolia Jacquin (1791: 215) from living material grown at the Imperial Gardens at Schönbrunn in Vienna or from the University Botanic Garden (Rix 1989), originally collected by Georg Scholl and Franz Boos in Namaqualand, South Africa. This led Baker (1873) to present the combination Whiteheadia bifolia (Jacquin 1791: 215) Baker (1873: 226), a name that has been in use for more than 130 years. Further information on the history, ecology, pollination biology and distribution of this species can be found in Manning et al. (2011). The presence of a coma of sterile bracts at the apex of the inflorescence in W. bifolia makes it similar to Eucomis, which explains Jacquin’s original generic classification (Jacquin 1791). However, W. bifolia differs from Eucomis by distinct morphological characters, such as the 2 opposite, prostrate, semisucculent, fragile leaves; the squat, leafy spike of greenish flowers surmounted by a crown or coma of sterile floral bracts; the hooked style; and the papery, winged capsules (cf. Manning et al. 2011). This represents a unique syndrome of morphological characters that allowed recognition of Whiteheadia as a monotypic genus for more than a century. Müller-Doblies & Müller-Doblies (1997) described a second species in Whiteheadia, W. etesionamibensis Müller- Doblies & Müller-Doblies (1997: 82) from south western Namibia, which differs from W. bifolia by distinct characters, such as the narrower, papillose-ciliate, non-succulent bracts (not entire, subsucculent, much larger bracts); the honey- scented, ± white, subcampanulate flowers (not shallow, “sour” smelling, green, subrotate flowers); the perigone fused at the base to form a cup shaped tube (not a subrotate tube); perigone segments 10−13 mm long (not only 6−8 mm long); filaments acuminate (not thickened), and smaller anthers (Müller-Doblies & Müller-Doblies 1997). Recent studies based on molecular data included Whiteheadia in the synonymy of Massonia (Manning et al. 2004, 2011, Manning & Goldblatt 2012, 2013), thus resulting in an expanded Massonia characterised by its paired, prostrate leaves and filaments that are fused at the base (Manning et al. 2004). Manning et al. (2011) stated: “DNA data indicate that this species [W. etesionamibensis] falls between Whiteheadia and Massonia, rather than grouping immediately with Whiteheadia, providing grounds for concluding that it shared a recent common ancestor with Massonia. Available evidence, therefore, suggests that it occupies an evolutionary position intermediate between the two genera, as a result of which we elected to treat Massonia rather more broadly than before to include Whiteheadia, rather than embrace the alternative solution of describing an additional monotypic genus for W. etesionamibensis”. However, the topology of the phylogenetic tree by Manning et al. (2004) does not agree with their own conclusions. A sample of Whiteheadia etesionamibensis is sister to a clade (69% bootstrap) comprising one sample of W. bifolia which is consecutively sister to a clade (98% bootstrap) fitting Massonia in its traditional sense. The later phylogenetic relationships were also corroborated by Wetschnig & Pfosser (2003) and Pfosser et al. (2003). Therefore, W. etesionamibensis cannot be considered as intermediate between Whiteheadia s.s. and Massonia, but sharing a common ancestor which gave rise to both later traditionally accepted genera. The phylogenetic relationships of Whiteheadia etesionamibensis, Whiteheadia s.s. and Massonia (Wetschnig & Pfosser 2003, Pfosser et al. 2003, Manning et al. 2004) agree with their floral morphology. Flowers of Whiteheadia s.s. resemble some species of Massonia, especially of the Massonia depressa Houttuyn (1780: 424) complex, based on the short and broad perigone-filaments tube and the broad triangular exposed ovary. Furthermore, the recently described Massonia obermeyerae Martínez-Azorín et al. (2015: 42) shows links to Whiteheadia s.s. based on strongly incurved filaments that are thickened and tapering to the tip, although they are again swollen in the region of the white and spongy anther connective and the fleshy unripe capsules that bear a long-lasting, fleshy, hooked style (Martínez- Azorín et al. 2015). The hypothesis that M. obermeyerae constitutes the link between Massonia and Whiteheadia (Obermeyer 1965, Martínez-Azorín et al. 2015) agrees with our phylogenetic studies (M. Martínez-Azorín and collaborators, in preparation), in which two samples of M. obermeyerae form a basal clade sister to all other Massonia species, suggesting that a common ancestor of this species gave rise to the genus Massonia. Our studies (not shown) also place several samples of W. bifolia as a monophyletic clade sister to the clade comprising M. obermeyerae plus the remaining Massonia species. However W. bifolia differs from all other Massonia species by its fragile and succulent leaves, and the elongated inflorescence with long, comose bracts and stout peduncle (Table 1), characters that support its acceptance as a genus in its own right. Wester et al. (2009) commented that “Whiteheadia bifolia has a specific combination of floral characters consistent with the rodent pollination syndrome (geoflory, visual inconspicuousness, bowl-shaped, robust flowers, easy accessible, very viscous nectar and a slightly sourish-nutty smell). In contrast, the second species of the genus Whiteheadia, the Namibian W. etesionamibensis U.Müll.-Doblies & D.Müll.-Doblies, has honey-scented, white flowers that suggest insect pollination (see Müller-Doblies & Müller-Doblies 1997)”, therefore underpinning the distinct floral differences between the two Whiteheadia species.

202 • Phytotaxa 221 (3) © 2015 Magnolia Press MARTÍNEZ-AZORÍN ET AL. FIGURE 1. Desertia etesionamibensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.Pinter & Wetschnig from the Dikwillem in Namibia, in cultivation in Graz, Austria on 15 December 2014 (corresponding to Müller-Doblies 88042C). a. Plant in flower; b. Inflorescence in apical view showing the small apical coma of sterile bracts. Scale bars 1 cm.

Manning et al. (2004) accepted Namophila urotepala Müller-Doblies & Müller-Doblies (1997: 77) as a monotypic genus based on morphological characters that are of similar degree or even weaker than those separating W. etesionamibensis and Massonia or Whiteheadia. The genus Namophila was segregated by Manning et al. (2004) and

Desertia, a new genus in Massonieae Phytotaxa 221 (3) © 2015 Magnolia Press • 203 Manning & Goldblatt (2013) from Massonia (including Whiteheadia) by the “Tepals caudate; capsule narrowly ovoid, not distinctly 3-lobed, remaining enclosed by the perianth at maturity” versus “Tepals acute but never caudate; capsule obtriangular and deeply 3-angled or winged, exposed from the withered perianth at maturity” in Massonia sensu lato. Our studies in Massonia reveal that several species of this latter genus do not produce deeply 3-angled or winged capsules, such as Massonia jasminiflora and related species that share obclavate gynoecium (Martínez-Azorín et al. 2013, 2014a, 2014b, Wetschnig et al. 2014). Therefore, the remaining morphological characters to accept Namophila versus Massonia are the caudate tepals, the capsule enclosed by the fleshy perigone at maturity and the narrow linear bracts (Table 1).

TABLE 1. Comparison of main diagnostic characters between Desertia, Whiteheadia, Massonia and Namophila. Desertia Whiteheadia Massonia Namophila Leaves Firm Fragile Firm Firm Inflorescence Long racemose, distinctly Long racemose, distinctly Subcapitate, not distinctly Subcapitate, not distinctly elongated above ground elongated above ground elongated above ground elongated above ground level level level level surmounted by a distinct surmounted by a distinct not surmounted by a distinct not surmounted by a distinct coma of small sterile floral coma of large sterile floral coma of sterile floral bracts coma of sterile floral bracts bracts bracts Bracts Lanceolate Ovate-triangular Lanceolate to widely ovate Narrowly linear Strongly papillose on both Glabrous, with entire or Glabrous or hairy with Glabrous with entire margin sides and margin minutely denticulate margin entire, denticulate or dentate margin Erect at early stages of Spreading or patent at early Erect to patent at early Erect at early stages of inflorescence development stages of inflorescence stages of inflorescence inflorescence development development development Flowers Perigone segments erect Perigone segments patent Perigone segments patent to Perigone segments patent from the perigone-filaments from the perigone-filaments strongly reflexed from the from the perigone-filaments tube, straight or slightly tube, curving upwards only perigone-filaments tube tube spreading at the upper portions Honey scented “Sour” scented Honey or “sour” scented “Sour” scented Style Straight Hooked Straight or hooked Straight Capsule Exposed from the withered Exposed from the fleshy Exposed from the withered Enclosed by the fleshy perigone at maturity perigone at maturity perigone at maturity perigone at maturity Seeds With sinuous rugosity Smooth Smooth or rugose Slightly rugose

Regarding Whiteheadia etesionamibensis, it shows subcampanulate white flowers, in which the tepals are more or less straight, suberect or slightly spreading, greatly differing from any species of Massonia, Namophila or Whiteheadia (Table 1), in which the free portion of tepals are patent to strongly reflexed, sometimes showing a sigmoid curve at the base. Only some species, among them W. bifolia, show tepals suberect in their upper portions, but they are patent from the mouth of the perigone tube and curve upwards again, usually due to the enclosing of the flowers by the large bracts. The inclusion of W. etesionamibensis in Massonia greatly compromises recognition of other genera in tribe Massonieae based on flower morphology. Therefore, we choose to accept W. etesionamibensis as a distinct genus. This solution is also supported by our phylogenetic studies (not shown). Müller-Doblies & Müller-Doblies (1997) described Whiteheadia etesionamibensis as having “perigonio subcampanulato albo” and flowers “± white” and the ovary “suddenly contracted into the [...] white style”. The description of the ovary fits an illustration of the type collection from the Aurus Mountains in Namibia (fig. 8g1 in Müller- Doblies & Müller-Doblies 1997: 81), in which an obovate ovary with truncate apex is shown. The cited illustration by Müller-Doblies & Müller-Doblies (1997) also compares flowers and bracts from the type locality with the collection Müller-Doblies 88141k from the Rooiberg. The ovaries of samples from the latter locality are narrower, subovate and slightly tapering to the style that is longer than that from the type locality. In the framework of a taxonomic revision of Massonia and related genera (Martínez-Azorín et al. 2013, 2014a, 2014b, 2015, Pinter et al. 2013, 2015, Wetschnig et al. 2012, 2014), we here describe a new genus, named Desertia, to accommodate Whiteheadia etesionamibensis. The new genus is based on distinct morphological, phylogenetic and biogeographic evidence as shown below. The study of living plants in cultivation from two populations of Desertia and further herbarium materials and images evidence

204 • Phytotaxa 221 (3) © 2015 Magnolia Press MARTÍNEZ-AZORÍN ET AL. that distinct morphological differences exist allowing the description of a second species in the genus, here named D. luteovirens, characterized by the yellow-greenish flowers, the ovate, narrower ovary, the shorter style, bigger capsules and the seed morphology. Furthermore, our phylogenetic data (not shown) also support differentiation of the newly described species, as commented below. A taxonomic treatment for Desertia, Namophila and Whiteheadia is also presented, including identification keys for all accepted taxa and complete morphological descriptions.

FIGURE 2. Desertia etesionamibensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.Pinter & Wetschnig from the Dikwillem in Namibia, in cultivation in Graz, Austria on 15 December 2014 (corresponding to Müller-Doblies 88042C). a. Inflorescence in lateral view, showing the small apical coma of sterile bracts; b. Flowers in apical (above) and lateral views (below); c. Bracts. Scale bars 1 cm.

Detailed morphological studies of species of Desertia, Massonia, Namophila and Whiteheadia from southern Namibia and the Northern Cape and Western Cape Provinces of South Africa were undertaken on natural populations and cultivated specimens as elaborated upon in Martínez-Azorín et al. (2007, 2009). Herbarium specimens from the herbaria ABH, CGE, G, GZU, GRA, L, M, MO, NBG, NU, NY, PRE, S, TCD, WAG, WIND, Z and ZSS (acronyms according to Thiers 2015) were studied. Orthography of geographical names and grid-number system follows Leistner & Morris (1976). Morphological measurements and illustrations of leaves and flowers were made on fresh and on herbarium material derived from wild plants. Author names of the cited taxa follow IPNI (2015).

Taxonomy of Desertia, Whiteheadia and Namophila

1. Desertia Mart.-Azorín, M.Pinter & Wetschnig gen. nov. (Figs. 1−4, 7−10; fig. 8 in Müller-Doblies & Müller- Doblies 1997: 81).

Desertia differs from Whiteheadia (W. bifolia) and Massonia by the papillose bracts and subcampanulate flowers with straight, suberect or slightly spreading free portion of tepals and the sinuous-rugose seeds (Table 1). Whiteheadia differs from Desertia by the larger, leafy, glabrous bracts, which are patent at early stages of inflorescence development; the subrotate flowers with patent free portions of tepals with regards to the perigone tube, which curve upwards in the upper portions; the larger anthers before dehiscence; the hooked style; and the smooth seeds (Table 1). Massonia differs from Desertia by the subcapitate, sessile, raceme, not distinctly elongated above ground level; the bracts lacking distinct papillae; the absence of a distinct apical coma of sterile bracts; the free portion of tepals being patent to strongly reflexed with regards to the perigone-filaments tube, at least at the base; and the smooth seeds (Table 1). Type (holotype):—Desertia etesionamibensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.Pinter & Wetschnig Decidous geophyte. Bulb solitary, hypogeal, ovate-globose. Leaves two, synanthous, opposite, appressed to the ground, green with undefined, longitudinal, darker green stripes. Inflorescence, spiciform-racemose, multiflowered, distinctly elongated above ground level, surmounted by a coma of small, sterile floral bracts. Bracts lanceolate, membranous, strongly papillose on both sides and margin, erect to connivent at early stages of inflorescence development. Flowers subcampanulate, white or yellowish-green, patent to suberect, shortly pedicellate, honey-scented; tepals glabrous and entire or slightly papillose and denticulate on margins, straight, suberect or slightly spreading, fused below to form a short tube; filaments adnate to perigone and arising above the perigone tube, uniseriate, shortly connate above perigone; anthers yellow with pink-violet flush, dorsifixed. Ovary green, ovate to obovate-oblong; style mostly white, narrow, erect or slightly arcuate; stigma minute. Capsule widely ovate to obovate in lateral view, trigonous and deeply winged in apical view. Seeds more or less globose, apiculate, with a distinct hilum and a slightly prominent raphe, with sinuous rugosity. Etymology:—Desertia (desertum lat. = desert); referring to desert habitats where the representatives of the new genus occur. Biogeography and ecology:—Endemic to south western Namibia and north western South Africa. It occurs from farm Weissenborn and Dikwillem in Namibia to Richtersveld in South Africa, found among rocks and boulders on rocky hillslopes.

1a. Desertia etesionamibensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.Pinter & Wetschnig comb. nov.

Bas.: Whiteheadia etesionamibensis Müller-Doblies & Müller-Doblies (1997: 82) (Figs. 1−4, 5a; figs. 8a, 8b, 8e, 8f, 8g1, 8i1, 8h, 8j in Müller-Doblies & Müller-Doblies 1997: 81). Type:—NAMIBIA. Witputz (2716): Aurus Mts., E of summit (Diamond area nº1) (-CB), S-facing slope, ca. 800 m of elevation, 18 September 1988, Müller-Doblies 88144n (holotype, WIND; isotypes, B, BTU, G, K, LI, M, MO, NBG, PRE, S, Z; not seen, apparently not yet deposited in any of the cited herbaria, including the holotype).

Deciduous geophyte. Bulb 2−3 × 1−2 cm, solitary, hypogeal, ovate-globose. Leaves 5−10 × 1.5−6 cm, two, synanthous, opposite, appressed to the ground, ovate, with entire to minutely denticulate margin, green, with undefined, longitudinal, darker stripes. Inflorescence 3−7 cm long, solitary, spiciform-racemose, distinctly elongated above ground level, with 10−30 flowers, surmounted by a coma of small, inconspicuous, sterile, floral bracts. Bracts 10−15 × 4−6 mm, lanceolate,

206 • Phytotaxa 221 (3) © 2015 Magnolia Press MARTÍNEZ-AZORÍN ET AL. FIGURE 3. Desertia etesionamibensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.Pinter & Wetschnig from the Dikwillem in Namibia, in cultivation in Graz, Austria on 15 December 2014 (corresponding to Müller-Doblies 88042C). a. Dissected bud and flower before anthesis in lateral view; b. Dissected flower at anthesis in lateral view; c. Gynoecia in lateral (above) and apical views (below). Scale bars 1 cm.

Desertia, a new genus in Massonieae Phytotaxa 221 (3) © 2015 Magnolia Press • 207 membranous, papillose on both sides and margin, the uppermost ca. 7 × 3 mm, white with a central longitudinal green band on the upper half, mostly hidden by flowers. Flowers subcampanulate, patent to suberect, pure white with a green longitudinal central band in the upper portion of tepals, honey-scented; tepals 10−16 × 3−4 mm, lanceolate, acute, glabrous, entire, straight, suberect or slightly spreading, fused below to form a short tube 3−4 mm long; filaments adnate to perigone and arising above the perigone-filaments tube, uniseriate, free portions 4−6 mm long, lanceolate, shortly connate above perigone for ca. 1 mm; anthers before dehiscence ca. 2 mm long, ca. 1 mm long after dehiscence. Ovary 4−5.5 × 3.5−4.5 mm, green, obovate-oblong; style 4−7 mm long, 0.6−1 mm wide at the base; mostly white, greenish in the lower third, narrow, erect or slightly arcuate, stigma minute. Capsule 7−11 × 7−13 mm, widely obovate in lateral view, deeply three lobed or winged in apical view. Seeds 2−2.3 × 1.5−1.8 mm, more or less globose, apiculate, with a distinct hilum and a slightly prominent raphe, glossy black, with distinct sinuous rugosity.

FIGURE 4. Desertia etesionamibensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.Pinter & Wetschnig from the Dikwillem in Namibia, in cultivation in Graz, Austria on 19 February 2014 (a) and 4 April 2014 (b,c) (corresponding to Müller-Doblies 88042C). a. Plant in fruit before capsule dehiscence; b. Infructescence with dehiscing capsules; c. Dehiscing capsules in lateral views. Scale bars 1 cm.

208 • Phytotaxa 221 (3) © 2015 Magnolia Press MARTÍNEZ-AZORÍN ET AL. FIGURE 5. Seed morphology of Desertia etesionamibensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.Pinter & Wetschnig (a), Desertia luteovirens Mart.-Azorín, M.Pinter & Wetschnig (b), Whiteheadia bifolia (Jacq.) Baker (c) and Namophila urotepala U.Müll.- Doblies & D.Müll.-Doblies (d). 1. Seed, lateral view; 2. Seed, raphal view. Scale bars 0.5 mm.

Desertia, a new genus in Massonieae Phytotaxa 221 (3) © 2015 Magnolia Press • 209 Comments on the type collection:—We were not able to study the type material of D. etesionamibensis, since it has not yet been deposited in any of the herbaria cited by Müller-Doblies & Müller-Doblies (1997), including the holotype at WIND (E. Strauss pers. comm.). However, figure 8 in Müller-Doblies & Müller-Doblies (1997: 81) includes detailed drawings of bracts, flowers and capsules from the type collection in comparison with a collection from the Rooiberg. Provided that Whiteheadia etesionamibensis was described as having white flowers, and that the illustration of the gynoecium from the type collection by Müller-Doblies & Müller-Doblies (1997) shows widely obovate and truncate ovary that extends into a shorter style with regards to the collection form the Rooiberg, point out that Desertia etesionamibensis, as originally described, represents the white-flowered species with broader, truncate ovaries and shorter styles among other characters, as accepted in the present study. Etymology:—The specific epithet means growing in winter-rainfall region characterised by ethesial winds, referring to etesiai winds that (in ancient Greece) used to bring winter rainfall; the term ‚ethesial regions’ is still used by climatologists to designate regions experiencing mediterranean-type climate (Müller-Doblies & Müller-Doblies 1997). Biology:—Desertia etesionamibensis flowers in the wild from May to July, and fruits arise from June to August. In cultivation in the Northern Hemisphere (Graz, Austria) this species flowers from late October to January, and capsules are well developed from February to April. Distribution:—Desertia etesionamibensis is known to us from four localities in south western Namibia, farm Weissenborn and Dikwillem in the Lüderitz district, the Aurus Mountains (the type locality) and the Hunsberge (Fig. 6).

FIGURE 6. Known distribution of Desertia etesionamibensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.Pinter & Wetschnig (red squares) and Desertia luteovirens Mart.-Azorín, M.Pinter & Wetschnig (blue circles) in southern Namibia and north western South Africa.

Habitat:—Among rocks and boulders on rocky hillslopes. Diagnostic characters:—Desertia etesionamibensis is characterized by the white flowers; the glabrous tepals with entire margin; the obovate and truncate ovary, 3.5−4.5 mm wide; the style 5−7 mm long; the capsules 7−11 × 7−13 mm; and the seeds glossy black, with a slightly prominent raphe and distinct sinuous rugosity. Additional material studied:—NAMIBIA. Aus (2616): SWA, Lüderitz district, farm Weissenborn, ravine near farmhouse (-AB), between rocks, 4 July 1949 [in flower and fruit], H. Kinges 2387 (M0274451!, PRE0049670!); Aus (2616): Dikwillem, ca. 20 km NW of Aus (-AC), rock crevices, ca. 900–1150 m elevation, ex hort. in Graz, Austria on 15 December 2014 [in flower], corresponding to Müller-Doblies 88042C (GZU!); Chamaites (2717): Nuobrivier (-CA), unweit Apollogrotte, Hunsberge, am Berghang im Gestein, 9 June 1976 [in fruit], W. Giess & M. Müller 14305 (M, WAG1158220!, WIND000003262 photo!).

1b. Desertia luteovirens Mart.-Azorín, M.Pinter & Wetschnig sp. nov. (Figs. 5b, 7−10; fig. 8d, 8g, 8g2, 8g3, 8i2, 8k in Müller-Doblies & Müller-Doblies 1997: 81).

210 • Phytotaxa 221 (3) © 2015 Magnolia Press MARTÍNEZ-AZORÍN ET AL. Desertia luteovirens is in some extent similar to its relative D. etesionamibensis, but it differs by the pale yellow-greenish flowers; the tepals slightly papillose with denticulate margins; the ovary ovate, tapering to the style, 2.5−3.2 mm wide; the style 3.5−4 mm long; the capsule 12−21 × 13−25 mm; and the seeds dull black, with an indistinct raphe and finely sinuous rugosity. Type:—NAMIBIA. Witputz (2716): ca. 15.5 km NW of Rosh Pinah (-DC), ca. 756 m of elevation, 19 February 2015 ex hort. in Austria, Steiermark, Graz, Botanical Garden of the Institute of Plant Sciences of the University of Graz, C. Craib s.n. (holotype, WIND! isotypes, ABH!, GZU!).

Deciduous geophyte. Bulb 2−4 × 1.5−3 cm, solitary, hypogeal, ovate-globose. Leaves 6−15 × 4−10 cm, two, synanthous, opposite, appressed to the ground, ovate, entire with minutely denticulate margin, green, with undefined, longitudinal, darker stripes. Inflorescence 7−11 cm long, solitary, spiciform-racemose, distinctly elongated above ground level, with 10−80 flowers, surmounted by a coma of small, sterile, linear-lanceolate floral bracts. Bracts 6−22 × 4−6 mm, lanceolate, membranous, densely papillose on both sides and margin, the uppermost ca. 7 × 2−3 mm, greenish with a central longitudinal darker green band, mostly hidden by flowers. Flowers subcampanulate, patent to suberect, greenish-yellow with green longitudinal bands in the upper portion of tepals, patent to suberect, honey-scented; tepals 10−16 × 3−4 mm, lanceolate, acute, slightly papillose with denticulate margins, straight, suberect or slightly spreading, fused below to form a tube 3−4 mm long; filaments adnate to perigone and arising above the perigone-filaments tube, uniseriate, free portions 4−6 mm long, lanceolate, shortly connate above perigone for ca. 1 mm; anthers before dehiscence ca. 2 mm long, ca. 1 mm after release of pollen. Ovary 4.5−5.5 × 2.5−3.2 mm, green, ovate, tapering to the style; style 3.5−4 mm long, 0.5−0.7 mm wide at base, mostly white, greenish in the lower third, narrow, erect; stigma minute. Capsule 12−21 × 13−25 mm, widely subovate to obovate in lateral view, trigonous and deeply winged in apical view. Seeds 1.9−2.1 × 1.8−1.9 mm, more or less globose, apiculate, with a distinct hilum and an indistinct raphe, dull black, with finely sinuous rugosity. Etymology:—The specific epithet comes from “luteo” = yellow and “virens” = green; referring to yellow-green colour of the flowers. Biology:—Desertia luteovirens flowers in the wild around January on the farm Spitskop in Namibia and in July in Richtersveld in South Africa. In cultivation in the Northern Hemisphere (Graz, Austria), D. luteovirens flowers from late January to April and capsules are well developed from March to May. It was observed that different plants from the type locality flowered at different times in successive years, and that flowering and fruiting plants occurred together. This may point out that the species is adapted to flower in response to suitable environmental conditions, adapting the anthesis to the rainfall periods. Under equal conditions of cultivation in Graz, Austria, D. luteovirens flowers later than D. etesionamibensis. Distribution:—The new species is known to us from several localities in south western Namibia, and north western South Africa. In Namibia it occurs in the Aurus Mountains, and the surroundings of the farm Spitskop, Rosh Pinah and Lorelei, whereas in South Africa it is only known from east of Sendelingsdrif in the Richtersveld National Park, the latter locality being the first record of Desertia in South Africa (cf. Manning & Goldblatt 2013) (Fig. 6). Habitat:—Among rocks and boulders on rocky hillslopes. Phylogenetic relationships:—Our phylogenetic studies on Desertia and related genera (not shown) evidence important differences between the two species of Desertia. Two samples of Desertia luteovirens showed a deletion of 94 base pairs in the trnL-F plastidial region, which is absent in Desertia etesionamibensis, this therefore adding further arguments for its recognition as a distinct species in the genus. Additional specimens and material studied:—NAMIBIA. Witputz (2716): Northern ridge of Aurus Mountains (-CB), mountain slope, stony/rocky soil, 683 m elevation, 11 August 2001 [in flower] C.A. Mannheimer 1592 (WIND77745.0 photo!; Mannheimer 2002: 68 photo!); Witputz (2716): Sperrgebiet, Aurus Mountains, lower peaks to SE of low neck SE of main beacon (-CB), granite kloof and slopes facing SW, 7 September 1992 [in flower] E.G.H. Oliver 10179 (WIND000059136 photo!); Witputz (2716): Distrikt Lüderitz—Süd, Numais-Bänke, Farm Spitskop (- DC), 1 January 1963 [in flower], H. Merxmüller & W. Giess 3408 (M0274455!); Witputz (2716): Lüderitz-Süd Distr., Rosh Pinah (-DD), mountain slopes above the mine, NE of the village, and flats below, rocks, 2 July 1947 [in bud], B. Nordenstam & J. Lundgren 491 (S09-7574!); Witputz (2716): Rosh Pinah, Farm Namuskluft, high mountain North of road, +- 5 km from house (-DD), shady, wet gorge, steep slope, in the cracks of a dark grey boulder, 589 m elevation, 10 September 2002, S. Bartsch & S. Loots SB1092 (WIND83150.0 photo!, Mannheimer et al. 2008: 29 photo!); Oranjemund (2816): N of Lorelei (-BB), growing on 35º slope with 0 aspect, 1 August 1989 [in flower and fruit], B. Strohbach 122 (WIND000056841 photo!). SOUTH AFRICA. Northern Cape. Oranjemund (2816): Near Swartpoort, Noorskoppe (referring to Euphorbia hottentota), Richtersveld National Park, high peaks just N of road (9 km from Sendelingsdrif on road to Oena) (-BB), succulent karroo, stony soil/rocky, 28 July 1993 [in flower], E.J. Van Jaarsveld 13247 (PRE763394.0!).

Desertia, a new genus in Massonieae Phytotaxa 221 (3) © 2015 Magnolia Press • 211 FIGURE 7. Desertia luteovirens Mart.-Azorín, M.Pinter & Wetschnig from ca. 15 km NW of Rosh Pinah in cultivation in Graz, Austria on 19 February 2015 (corresponding to the holotype Craib s.n.). a. Plant in flower; b. Inflorescence in apical view showing the small coma of sterile bracts. Scale bars 1 cm.

212 • Phytotaxa 221 (3) © 2015 Magnolia Press MARTÍNEZ-AZORÍN ET AL. FIGURE 8. Desertia luteovirens Mart.-Azorín, M.Pinter & Wetschnig from ca. 15 km NW of Rosh Pinah in cultivation in Graz, Austria on 19 February 2015 (corresponding to the holotype Craib s.n.). a. Dissected inflorescence in lateral view; b. Flowers in apical (above) and lateral views (below); c. Bracts. Scale bars 1 cm.

Desertia, a new genus in Massonieae Phytotaxa 221 (3) © 2015 Magnolia Press • 213 FIGURE 9. Desertia luteovirens Mart.-Azorín, M.Pinter & Wetschnig from ca. 15 km NW of Rosh Pinah in cultivation in Graz, Austria on 19 February 2015 (corresponding to the holotype Craib s.n.). a. Dissected bud and flower before anthesis in lateral view; b. Dissected flower at anthesis in lateral view; c. Gynoecia in lateral (above) and apical views (below). Scale bars 1 cm.

214 • Phytotaxa 221 (3) © 2015 Magnolia Press MARTÍNEZ-AZORÍN ET AL. Other specimens of Desertia cited in Müller-Doblies & Müller-Doblies (1997) but not available:—NAMIBIA. Aus (2616): Dikwillem, ca. 20 km NW of Aus (-AC), rock crevices, ca. 900–1150 m elevation, 27 July 1988, Müller- Doblies 88042C (B, BTU, K, LI, PRE, WIND); Witputz (2716): Rooiberg, top and upper S slope (Diamond area nº1) (-CB), stony ground and crevices, ca. 1050 m elevation, 16 September 1988, Müller-Doblies 88141k (BTU, WIND); Witputz (2716): Numaeis, südl. Witputz, (-DC), September 1957, Rusch f. 4688 (M, on loan to U. & D. Müller- Doblies); Witputz (2716): Spitskop, valley with waterfall, 6 km from Rosh Pinah road, kloof around the seasonal waterfall (-DC), ca. 700 m elevation, 6 August 1988 in bud, Müller-Doblies 88064b (BTU); Witputz (2716): farm Spitskop, valley entry 1,2 km S of farm house (-DC), rocky S facing slope, ca. 650 m elevation, 27 September 1989 nearly flowering, Müller-Doblies 89123g (B, BTU, NBG, WIND, S, Z); Witputz (2716): Namuskloof, 7 km N of farm house (-DD), S facing slope with quartzite, ca. 450 m elevation, 6 August 1988, Müller-Doblies 88062e (BTU, WIND); Oranjemund (2816): Diamond area nº1, Obib Mountain Peak, (-BA), rocky S facing slope, 3 September 1989, A.E. Van Wyk 9005 (PRE, PRU, WIND).

FIGURE 10. Desertia luteovirens Mart.-Azorín, M.Pinter & Wetschnig from ca. 15 km NW of Rosh Pinah in cultivation in Graz, Austria on 18 March 2014 (a), 02 May 2014 (b) (corresponding to Craib s.n.). a. Plant in fruit before capsule dehiscence; b. Infructescence with dehiscing capsules; c. Dehiscing capsule in lateral view. Scale bars 1 cm.

1. Flowers white; tepals glabrous, entire; ovary 3.5−4.5 mm wide, obovate, suddenly narrowed into the style; style 4−7 mm long; capsule 7−11 × 7−13 mm; seeds glossy black, with a slightly prominent raphe and distinct sinuous rugosity ...... 1a. D. etesionamibensis - Flowers pale yellow-greenish; tepals slightly papillose with denticulate margins; ovary 2.5−3.2 mm wide, ovate, tapering to the style; style 3.5−4 mm long; capsule 12−21 × 13−25 mm; seeds dull black, with an indistinct raphe and finely sinuous rugosity ...... 1b. D. luteovirens

2. Whiteheadia bifolia (Jacq.) Baker (1873: 226) ≡ Eucomis bifolia Jacquin (1791: 215) ≡ Basilea bifolia (Jacq.) Poiret (1811: 591) ≡ Massonia bifolia (Jacq.) J.C.Manning & Goldblatt in Manning et al. (2004: 564) (Figs. 11−12). Type (lectotype, designated by Jessop 1976: 433):—SOUTH AFRICA. without precise locality or collector (t. 449 in Jacquin 1791).

= Melanthium massoniaefolium Andrews (1804: t. 386). Type (lectotype, designated by Manning et al. 2011: 331):—SOUTH AFRICA. without precise locality, unknown collector (t. 368 in Andrews 1804). = Whiteheadia latifolia Harvey (1868: 396). Type:—SOUTH AFRICA. Namaqualand, Modderfontein, Whitehead s.n. (holotype, TCD!).

Deciduous geophyte. Bulb 1−2.5 × 1−3 cm, solitary, hypogeal, ovate-globose, with papery, brownish outer tunics. Leaves 8−24 × 5−19 cm, two, synanthous, opposite, appressed to the ground, ovate to suborbicular, pale green, fragile and somewhat succulent, glabrous, smooth, with depressed longitudinal nerves and entire margin. Inflorescence 5−18 cm long, solitary, spiciform-racemose, distinctly elongated above ground level, with 10−50 flowers, surmounted by a distinct coma of sterile, ovate-lanceolate floral bracts; peduncle 2−6 cm long; axis of the inflorescence stout, distinctly swollen, tapering below. Bracts 15−50 × 8−25 mm, large, leafy, green, ovate-triangular, acute, glabrous, with entire to minutely denticulate margin, patent in the lower half, suberect to slightly reflexed at the apex, concave below to enclose the flowers, persistent in fruit, patent at early stages of development of the inflorescence. Flowers suberect, pale green, “sour” scented; tepals ovate, glabrous, entire, fused at the base to form a subrotate perigone-filaments tube of ca. 4 mm long, the free portion of tepals 7−9 × 4−6 mm, patent from the tube but curving upwards mostly due to the enclosing by the bracts; filaments pale green, adnate to perigone and arising above the perigone-filaments tube, uniseriate, free portions 6−9 mm long, shortly connate above perigone for ca. 1 mm long, distinctly arcuate-incurved towards the style, swollen and triangular below, tapering up to the apex, but slightly thickening again near the anther junction with a distinct white zone; anthers pale yellow, before dehiscence ca. 5 mm long, ca. 3 mm after dehiscence. Ovary 5−6.5 × 5−7 mm, green, subpentagonal; style green, erect but hooked at the tip, 6−9 mm long; stigma minute. Capsule 1.9−2.6 × 1.8−2.8 cm, broadly obovoid-globose or obtriangular, three-winged, papery. Seeds 1.5−2.4 × 1.3−1.8 mm, more or less globose, apiculate, with a distinct hilum, glossy black, smooth. Biology:—Whiteheadia bifolia flowers in the wild from May to October and fruits are developed from June to December. In cultivation in the northern hemisphere (Graz, Austria) this species flowers from November to January, and capsules are well developed from late December to April. This species is pollinated by rodents (Wester et al. 2009, Wester 2009a, 2009b, 2010, 2011, 2015). Distribution:—Whiteheadia bifolia occurs from the Karasburg area in southern Namibia to Namaqualand as far south as the Cederberg Mts. in South Africa. Habitat:—Whiteheadia bifolia grows in sheltered and shaded places between rocks and boulders in thin soils rich in nutrients originating from animal middlens of for instance Namaqua Rock Mice and Rock Elephant Shrews. Taxonomic relationships:—Whiteheadia bifolia is characterized by the long racemose-spiciform inflorescence, clearly elongated above ground level, with large leafy bracts that are longer than the flowers, topped by a coma of large sterile floral bracts; bracts glabrous, entire to minutely denticulate on margins, subsucculent, patent at early stages of inflorescence development; flowers subrotate with shallow, “sour” smelling; perigone segments patent from the perigone-filaments tube, curving upwards only at the upper portions; anthers ca. 4 mm long before dehiscence; style hooked; and leaves, semisucculent and fragile. Some species of Massonia share similar flower morphology with W. bifolia, but all known Massonia species differ from Whiteheadia by the inflorescence subcapitate, not clearly elongated above ground level, and not surmounted by a distinct coma of large sterile floral bracts, and the non-fragile or subsucculent leaves (Table 1). Desertia differ from Whiteheadia by the bracts strongly papillose on both sides and margin, that are erect at early stages of inflorescence development; the flowers subcampanulate, honey-scented; perigone segments erect from the perigone-filaments tube, straight or slightly spreading; anthers ca. 2 mm long before dehiscence; style straight; and leaves not succulent or fragile (Table 1).

216 • Phytotaxa 221 (3) © 2015 Magnolia Press MARTÍNEZ-AZORÍN ET AL. FIGURE 11. Whiteheadia bifolia (Jacq.) Baker from Narap, Northern Cape, South Africa in cultivation in Graz, Austria on 7 December 2013 (corresponding to WW04445). a. Plant in flower; b. Bract of the apical coma; c. Bract of flowers; d. bud, flower and dissected flower in lateral (above) and apical views (below). Scale bars 1 cm.

Desertia, a new genus in Massonieae Phytotaxa 221 (3) © 2015 Magnolia Press • 217 FIGURE 12. Whiteheadia bifolia (Jacq.) Baker from Narap, Northern Cape, South Africa in cultivation in Graz, Austria on 7 December 2013 (corresponding to WW04445). a. Flower with fleshy capsule in lateral view; b. Fleshy capsule in apical view; c. Dehiscing capsule in lateral view; d. Dehiscing capsule in apical view showing the seeds. Scale bars 1 cm.

Additional material studied:—NAMIBIA. Vioolsdrift (2817): Gamkarivier, 5 km above river mouth (-AD), S- facing steep slope, 200 m s.m., 9 August 1988, Müller-Doblies 88073c (BTU!, cited in Müller-Doblies & Müller- Doblies 1997); Warmbad (2818): District Warmbad, Farm Sperlingspütz: WAR259 (-CA), auf Gesteinskuppe mit Pachypodium namaquanum, 15 May 1963 [in leaves with sprouting inflorescence], W. Giess, O.H. Volk & B. Bleissner 6972 (PRE0049671!, M0274453!, WIND00003259 photo!); Warmbad (2818): Warmbad, Farm Sperlingspütz: WAR259 (-CA), zwischen Felsen, 27 May 1972 [starting to develop the inflorescence], W. Giess & M. Müller 12248 (PRE0239457!, M0274454!, WIND photo!); Witputz (2716): Namibia, Sud Witputz, N of Farmhouse (-DA), 1989, Lavranos s.n. (ZSS!); SOUTH AFRICA. Vioolsdrif (2817): Richtersveld, Rosyntjiesberg, neck north of Lelieshoek (-AC), karoo vegetation, well-drained stony, clay loam, S aspect, 30 August 1977 [in fruit], Oliver, Tölken & Venter

218 • Phytotaxa 221 (3) © 2015 Magnolia Press MARTÍNEZ-AZORÍN ET AL. 336 (PRE648200!); Vioolsdrif (2817): Richtersveld (-AC), ca. 460 m elevation, 29 May 2014, Pieter van Wyk (photo! iSpotnature: http://www.ispotnature.org/es/node/587535?nav=related); Springbok (2917): ca. 6 km north of R382 to Lekkersing (-AC), quarzite mountain, in full shade on the eastern face of a cliff, ca. 278 m elevation, 08 August 2012 [in fruit], Pieter van Wyk (photo! iSpotnature: http://www.ispotnature.org/es/node/514085?nav=related); Springbok (2917): at turn off 8.5 km W of Steinkopf, 5 km on road to Kosies (-BA), stony ground, ca. 1000 m s.m., 8 August 1979, Müller-Doblies 79167e (BTU, cited in Müller-Doblies & Müller-Doblies 1997); Springbok (2917): near Klipfontein (-BA), Bolus 6565 (BOL, G 00420204!); Springbok (2917): Steinkopf, ca. 4 km NE of new petrol station (-BB), gravelly ground, 11 August 1988, Müller-Doblies 88078e (BTU, cited in Müller-Doblies & Müller-Doblies 1997); Springbok (2917): on rocky hills near Okiep (-DB) & Steinkopf (-BC), August 1925 [in flower], R. Marloth 6766 (PRE 0049675!); Springbok (2917): Ratelpoort, north of Springbok (-BD), G.N. 19225, 12 October 1961 [in flower], D.S. Hardy 542 (PRE0049677!, M0274457!); Springbok (2917): Ratelpoort, north of Springbok (-BD), 23 August 2011 [in fruit], M. Martínez-Azorín s.n. (ABH Photo!); Springbok (2917): Ratelpoort, 25 km N of Springbok on road N7 (-BD), quartzite ridge, 800–900 m s.m., 8 August 1979, Müller-Doblies 79163f (BTU!, cited in Müller- Doblies & Müller-Doblies 1997); Springbok (2917): About 10 km from Nababeep on the pipeline road to Spektakelberg (-DA), 5 August 1974, P. Goldblatt 2261 (MO2258364!, S15-24586!, WAG1158222!); Springbok (2917): 1 kilometer from Nababiep on road to Windhoek-Springbok highway (-DB), on rocky, boulder strewn hillside, 305 m elevation, 25 August 1987 [in fruit], A. Nicholas 2475 (PRE728944!); Springbok (2917): ca. 1.5 km north of Concordia (-DB), 2 October 2010 [in fruit], M. Martínez-Azorín s.n. (ABH Photo!); Springbok (2917): ca 1 km west of Concordia to Orbicular kopie (-DB), 1013 m elevation, 28 August 2011 [in fruit], M. Martínez-Azorín s.n. (ABH Photo!); Springbok (2917): Mesklip (-DD), Lewis 1396 (SAM, cited in Jessop 1976); Gamoep (2918): Aggeneys (-AA), ca. 697 m elevation, 9 August 2011 [in fruit], Francois Kriel (photo! iSpotnature: http://www.ispotnature.org/es/node/ 531080?nav=related); Gamoep (2918): Aggeneys (-BB), G nº.20218 (SJ), July 1975 [in flower], D.S. Hardy 3836 (PRE0519877!); Gamoep (2918): Aggeneys (-BB), ca. 1020 m elevation, 20 April 2014 [sprouting inflorescence], Francois Kriel (photo! iSpotnature: http://www.ispotnature.org/es/node/581320?nav=related); Gamoep (2918): Bushmanland, Aggenys Mountain, west of Pofadder, Soutkloof area WSW of Aggenys farm (-BB), stony well-drained, clay loam soil, 24 August 1977 [in fruit], Oliver, Tölken & Venter 56 (PRE648199!); Gamoep (2918): Little Namaqualand, Granitkuppe 17 Meilen von Springbock nach Poffadder (-CA), zwischen Felsen, 13 September 1963, H. Merxmüller & W. Giess 3784 (M0274458!); Gamoep (2918): Ou Areb se Berge, 36 km ENE of Springbok on Pofadder road (-CA), quartzite ridge, ca. 1000 m s.m., 18 August 1988, Müller-Doblies 88109b and 88163b (BTU, cited in Müller-Doblies & Müller-Doblies 1997); Pofadder (2919): Pella se Berge, steep mountain slope below south-facing large cliffs east of Piet se Kloof (-AA), ca. 1000 m elevation, 18 June 2006 [in fruit] C. Craib s.n. (photo GZU!); Hondeklipbaai (3017): Northern Cape, Namaqualand district, Namaqua National Park, between Skilpad & Soebatsfontein (-BB), granite, 555 m elevation, 15 August 2005 [in flower], H.M. Steyn 717 (PRE752038.0!); Hondeklipbaai (3017): Kersboschfontein, SW slopes of Brandberg (-BB), 2 October 1989, C.Bayer & Müller-Doblies 89140y (BTU, cited in Müller-Doblies & Müller-Doblies 1997); Hondeklipbaai (3017): Kamieskroon (-BB), Thorne s.n. sub SAM 48856 (SAM, cited in Jessop 1976); Hondeklipbaai (3017): Kamieskroon (-BB), 8 July 2012 [in flower], Etwin Aslander (photo! iSpotnature: http:// www.ispotnature.org/es/node/502979?nav=related); Hondeklipbaai (3017): ca. 18 km north of Kamieskroon, (-BB), 30 August 2009 [in fruit], W. Wetschnig 03959 (photo!); Hondeklipbaai (3017): 11 miles WSW of Garies (-DB), Acocks 14926 (PRE, cited in Jessop 1976); Kamiesberg (3018): Kamiesberg, Studers Pass, 13,7 miles from Garies to Kamiesberg (-AC), dry W slope, granite rocks, 2000 feet elevation, 27 August 1967 [in fruit], M.F. Thompson 425 (PRE0049679!); Kamiesberg (3018): area of Rooimuskop, hill N of Loeriesfontein/Kliprand road (-DB), 12 km E of Vanrhynsdorp/Pofadder road, 11 August 1980, Müller-Doblies 88079b (BTU, cited in Müller-Doblies & Müller- Doblies 1997); Kamiesberg (3018): Kamieskroon, Paardehoek, 700 m elevation, 3 September 2001 [in flower], J. Venter 9579 (MO5438478!, NY00210984!); Vanrhynsdorp (3118): Heerenlogement (-DB), rocky slopes 400–500 m s.m., 05 November 1978, V. Müller-Doblies 78063m (BTU, cited in Müller-Doblies & Müller-Doblies 1997); Vanrhynsdorp (3118): Doringriver bridge, on old Klawer/Clanwilliam road (-DB), rock crevices, 14 March 1979, Müller-Doblies 79124c (BTU, cited in Müller-Doblies & Müller-Doblies 1997); Vanrhynsdorp (3118): Giftberg, Compton 20842 (NBG, cited in Jessop 1976); Vanrhynsdorp (3118): Gifberg (-DB), ca. 930 m elevation, 11 August 2014 [in fruit], Lynn McMaster (photo! iSpotnature: http://www.ispotnature.org/es/node/600834?nav=related); Calvinia (3119): Bokkeveld, near Nieuwoudtville (-AC), 900 m elevation, October 1916 [in flower], R. Marloth 8343 (0049673!, 0049674!); Calvinia (3119): Doringbos, S-facing rock ledges at the Doring River (-AC), 1 September 1980, Müller-Doblies 80130w (BTU, cited in Müller-Doblies & Müller-Doblies 1997); Calvinia (3119): Bokkeveld plateau, Papkuilsfontein, ca. 22 km S of Nieuwoudtville (-CA), road toward waterfall viewing area, 25 August 2009 [in fruit], C. Davidson 11541 (MO6320984!); Calvinia (3119): Nieuwoudtville, Oorlogskloof Nature Reserve (-CA),

Desertia, a new genus in Massonieae Phytotaxa 221 (3) © 2015 Magnolia Press • 219 702 m elevation, 17 August 2001, W.A.J. Pretorius 742 (NBG181402.0, MO5902380!); Calvinia (3119): Botterkloof, top of the pass (-CD), TMS rock crevices, 01 September 1980, Müller-Doblies 80127ab (BTU, cited in Müller-Doblies & Müller-Doblies 1997); Calvinia (3119): top of Botterkloof (-CD), Esterhuysen 3871 (BOL, cited in Jessop 1976); Calvinia (3119): Botterkloof (-CD), 17 August 2014 [in fruit], ca. 626 m elevation, Chris (photo! iSpotnature: http:// www.ispotnature.org/es/node/601697?nav=related); Clanwilliam (3218): Kransvleibergen, Leipoldt 322 (SAM, cited in Jessop 1976); Clanwilliam (3218): Kliphuis Trail midway start to cross roads (-BB), ca. 976 m, 17 September 2011 [in fruit], Anthony Rebelo (photo! iSpotnature: http://www.ispotnature.org/es/node/468557?nav=related); Wuppertal (3219): Pakhuisberg (-AA), in saxosis, 2000 feet elevation, 10 August 1897, Schlechter 10801 (Z000087975!); Wuppertal (3219): Pakhuis, Esterhuysen 3157 (BOL, cited in Jessop 1976); Wuppertal (3219): Wolfberg, Algeria (- AC), ca. 800 m elevation, 14 July 2012 [in flower], Patrick Lane (photo! iSpotnature: http://www.ispotnature.org/es/ node/483269?nav=related); Langebergen, Platklip (Granite), 09 September 1926 [in flower], R. Marloth 12884 (PRE0049676!).

FIGURE 13. Known distribution of Whiteheadia bifolia (Jacq.) Baker (red circles) and Namophila urotepala U.Müll.-Doblies & D.Müll.- Doblies (blue triangles) in southern Namibia and western South Africa.

3. Namophila urotepala Müller-Doblies & Müller-Doblies (1997: 77) (Figs. 14−15; fig. 6 in Müller-Doblies & Müller- Doblies 1997: 78). Type:—NAMIBIA. Witputz (2716): farm Spitskop, valley with waterfall, 6 km from Rosh Pinah road (-DC), kloof around the seasonal waterfall, ca. 700 m elevation, 6 August 1988 pfl., Müller-Doblies 88064g (Holotypus: WIND; isotypi: B, BOL, BR, BTU, E, G, K, LI, M, MO, NBG, PRE, S, Z; not seen, apparently not yet deposited in any of the cited herbaria, including the holotype).

Deciduous geophyte. Bulb 1.5−3 × 1.5−3 cm, solitary, hypogeal, ovate-globose, with papery brown outer tunics. Leaves 9−22 × 7−13 cm, two, synanthous, opposite, appressed to the ground, ovate to suborbicular, smooth, green with undefined darker green longitudinal bands, subsucculent, with entire margin. Inflorescence 2−4 cm long, solitary, subcapitate-racemose or shortly corymbose, not distinctly elongated above ground level, with 2−20 flowers, not surmounted by a distinct coma of sterile bracts; peduncle hypogeal, 1.5−4 cm long. Bracts 15−30 × 2−3.5 mm, narrowly linear, green, membranous, glabrous, inconspicuous and mostly hidden by flowers. Pedicels 7−15 mm long. Flowers up to 35 mm in diameter, stellate, greenish, “sour” scented; tepals narrowly ovate-lanceolate, 19−23 mm long, acute, glabrous, entire, green and persistent until capsule dehiscence, fused at the base to form a perigone-filaments tube of 4−6 ×4−6 mm, the free portion of tepals 13−16 mm long, 3.5−4.5 mm wide at base, with a caudate apex of 1−4.5 mm long, patent to slightly curving upwards; filaments white with a rose colour above, lanceolate, adnate to perigone and arising above the perigone-filaments tube, uniseriate, free portions 6−8 mm long, acuminate, shortly connate above perigone for ca. 2 mm, slightly incurved; anthers pale yellow, before dehiscence 3−4.5 mm long, ca. 2.5 mm long after dehiscence; pollen yellow. Ovary 4−6 × 3−4 mm, green, ovate to subconical, rounded in cross section, tapering to the style; style green with white apex, erect or slightly arcuate, 8−10 mm long; stigma minute. Capsule 7−10 × 6−7 mm, ovoid, papery, enclosed by the fleshy perigone at maturity. Seeds 1.5−2 × 1.2−1.8 mm, more or less globose, but distinctly apiculate, glossy black, finely rugose.

220 • Phytotaxa 221 (3) © 2015 Magnolia Press MARTÍNEZ-AZORÍN ET AL. FIGURE 14. Namophila urotepala U.Müll.-Doblies & D.Müll.-Doblies from farm Spitskop in Namibia (corresponding to the type collection Müller-Doblies 88064g = cult. number 7098-5) in cultivation in Graz, Austria on 21 November 2013. a. Plant in flower; b. Flowers in lateral (above) and apical views (below); c. Bracts; d. Dissected flower in lateral view. Scale bars 1 cm.

Desertia, a new genus in Massonieae Phytotaxa 221 (3) © 2015 Magnolia Press • 221 FIGURE 15. Namophila urotepala U.Müll.-Doblies & D.Müll.-Doblies from farm Spitskop in Namibia (corresponding to the type collection Müller-Doblies 88064 = cult. number 7098-5) in cultivation in Graz, Austria on 21 November 2013 (a, b) and 10 January 2013 (c, d). a. Gynoecia in lateral view; b. Flower with fleshy capsule in apical view; c. Flower with mature capsule in apical view; d. Flower with mature capsule in lateral view. Scale bars 1 cm.

Etymology:—The specific epithet refers to the caudate tepals. Biology:—Namophila urotepala flowers in the wild from June to September. In cultivation in the Northern Hemisphere (Graz, Austria) this species flowers from late October to January, and capsules are well developed from late February to April. Distribution:—Namophila urotepala was described from few localities in a single valley in south western Namibia, near the farm Spitskop, northwest of Rosh Pinah (Müller-Doblies & Müller-Doblies 1997). The study of a herbarium collection by E. Van Jaarsveld in July 2007 (Van Jaarsveld 21149 WIND!) allowed us to extend the distribution range of this species to the Sonneberg in the Hunsberge (Fig. 13).

222 • Phytotaxa 221 (3) © 2015 Magnolia Press MARTÍNEZ-AZORÍN ET AL. Habitat:—Namophila urotepala grows in sheltered sites on rocky hillslopes among rocks and boulders in mountains of arid, winter-rainfall area of southern Namibia. Taxonomic relationships:—Namophila urotepala is characterized by the subcapitate inflorescence, not clearly elongated above ground level, not surmounted by a distinct coma of sterile floral bracts; bracts narrowly linear; tepals distinctly caudate; and capsules enclosed by the fleshy perigone at maturity. Massonia differs from Namophila by the wider, lanceolate-ovate bracts, the tepals not distinctly caudate and the capsules exposed from the withered the perigone at maturity (Table 1). Additional materials studied:—NAMIBIA. Witputz (2716): Farm Spitskop, LU111 (-DC), Bergschlucht, SO- Berge, am Hang haüfig, meist zwischen Felsen, 15 June 1976 [in flower], W. Giess & M. Müller 14421 (M0274456!, MO2631840!, PRE0488875 photo!, WAG1158221!); Witputz (2716): Farm Spitskop, LU111 (-DC), Schlucht in Bergen im SO der Farm, saures Lavagestein, kleine Fläche unter Überhängen der Felswand, 25 September 1977 [in flower], M. Müller & W. Giess 32265 (M0223032!); Witputz (2716): Rosh Pinah, Farm Spitskop, entrance to waterfall valley, SW slope of Numaisspitze (-DC), on shaded S-facing slopes among boulders and scrub, 8 August 2000 [in flower], P. Goldblatt & J.C. Manning 11350 (NBG!); Witputz (2716): Rosh Pinah, north west of beacon, Sonberg (-DD), in crevices on cliffs, 600−900 m elevation, 3 September 2000 [in flower], P.V. Bruyns 8851 (NBG195868!); Chamaites (2717): Southern Namibia, Sonneberg, upper S slope, lower southern outlier of the Hunsberge (-CC), well drained rocky, loam soil, on S-facing dolomite mountain, 3 July 2007 [in flower], E. Van Jaarsveld 21149 (WIND000093340 photo!).

Identification key for Desertia and related genera

1. Inflorescence long racemose, clearly elongated above ground level, surmounted by a distinct coma of sterile floral bracts ...... 2 − Inflorescence subcapitate, not clearly elongated above ground level, not surmounted by a distinct coma of sterile floral bracts.....3 2. Bracts strongly papillose on both sides and margin; flowers subcampanulate; perigone segments erect or slightly spreading from the perigone-filaments tube; style straight ...... 1. Desertia − Bracts glabrous, entire to minutely denticulate on margins; flowers subrotate; perigone segments patent from the perigone-filaments tube, curving upwards only at the upper portions; style hooked ...... 2. Whiteheadia 3. Tepals distinctly caudate; bracts narrowly linear; capsule enclosed by the fleshy perigone at maturity ...... 3. Namophila − Tepals not distinctly caudate; bracts lanceolate to widely ovate; capsule exposed from the withered perigone at maturity ...... Massonia (not included here)

Acknowledgements

This work was partly supported by Fundación Ramón Areces (Spain), University of Alicante (Spain) and Karl-Franzens- University (Austria). Rhodes University (Dept. of Botany) and the Selmar Schonland Herbarium (GRA) provided working facilities to the first author between 2009 and 2011. The Department of Environment and Nature Conservation of Northern Cape Province and CapeNature of Western Cape Province provided permission to collect herbarium specimens to the first author (collecting permits numbers FLORA046/2010, FLORA047/2010, FLORA069/2011, FLORA070/2011 and AAA008-00031-0028 respectively). We thank the late C. Craib for sending seeds of Desertia luteovirens from the type locality. U. & D. Müller-Doblies kindly provided bulbs of Desertia etesionamibensis and Namophila urotepala. The National Herbarium of Namibia (WIND), and E. Strauss, C. Mannheimer and S. Rügheimer are thanked for sending interesting comments and for sharing photographs and locality details of Desertia. Special thanks go to S. Rügheimer for her valuable help checking morphological features of the herbarium collections at WIND. We also thank D. Bellstedt and L. Mucina for their help on our field trip in 2009. We acknowledge the help of all herbarium curators who kindly provided material and information.

References

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