Notes on the Strumariinae ( Amaryllidaceae-Amaryllideae ) . Six new taxa in Strumaria and Hessea from the central and northwestern Cape , South Africa , and southern Namibia

Newly described are four species and a subspecies of Strumaria and one species of Hessea. S. aestivalis Snijman from the Langberg and S. perryae Snijman from the Bokkeveld escarpment are rare species closely allied to S. pubescens W.F. Barker. S. discifera Marloth ex Snijman is widespread on the Bokkeveld and Roggeveld escarpments but S. discifera subsp. bulbifera Snijman which comprises several clonal populations, is narrowly restricted to the dolerite ridges near Nieuwoudtville. S. villosa Snijman, a rare species, is localised on quartz hills near Kosies in the Richtersveld. H. speciosa Snijman occurs in red sand and friable loam from southern Namibia to the central Cape.


INTRODUCTION
The Strumariinae, an exclusively southern African sub tribe of the Amaryllideae, is centred in the semi-arid winter rainfall region of the Cape Province.The often insignificant, hysteranthous leaves and short-lived autumn flowers of the species, are phenological characteristics which render many members of the subtribe insufficiently collected.Thus since the last review of the Strumariinae (Miiller-Doblies 1985) some 12 additional new taxa have been discovered, of which five have already been published (Snijman 1989;Snijman 1991).
The 37 known species of Strumariinae are currently placed in eight genera (Namaquanula D. & U. Miiller-Doblies, Kamiesbergia Snijman, Hessea Herb., Carpolyza Salisb., Strumaria Jacq., Bokkeveldia D. & U. Miiller-Doblies, Gemmaria Salisb.and Tedingea D. & U. Miiller-Doblies).Phylogenetic studies in the Strumariinae using cladistic analyses (Snijman in prep.) have shown that Strumaria, Bokkeveldia and Gemmaria are weakly defined and paraphyletic and that they are best treated as a single genus Strumaria.Although the necessary generic redelimi tation will be explained and effected elsewhere, it is important, notably for conservation purposes, to validate the names of the undescribed species of the subtribe.Four of the new species described here are assigned to Strumaria, here defined according to Ker-Gawler (1814), Bolus (1923), Barker (1943Barker ( , 1944)).The fifth new species is placed in Hessea sensu Miiller-Doblies (1985), which has proven to be a monophyletic genus with the exclusion of the poorly known species H. spiralis Baker.

1.
S. aestivalis Snijman, sp.nov., quoad tunicam luteam bulbi, folia pubescentia et flores infiindibuliformes ad S. pubescentem W.F. Barker accedit, sed ab ea concavitatibus latis inter filamenta interiora et stylum differt.Bulb solitary or occasionally forming bulblets, ovoid, 20-40 mm diam., with the outer fibrous covering ranging from brown to cream-coloured, fleshy and yellowish within; neck up to 70 mm long, rarely absent.Leaves absent at anthesis, 2 or rarely 3, recurved, lorate, 8 0 -280 x 15-26 mm.canaliculate, both surfaces densely pubescent with 2 mm long, patent, silky, white hairs; amplexicaul cataphyll shortly exserted, tipped with red, soon withering down; non-amplexicaul prophyll hidden in the bulb.Inflorescence widely spreading.60-100 mm across; scape 60-100 x 2 .5-4.0 mm.pale green to glaucous, sometimes flushed with pink, pubescent or glabrous, breaking off at the base in fruit; spathe valves  Distribution and phenology: the northwestern foothills of the Langberg, northwest of Loeriesfontein, is the only known locality of S. aestivalis (Figure 2).The population is confined to the southeast-facing banks of a seasonal stream, where the bulbs are aggregated in the shade of rocks or low shrubs, amongst shale chips overlying heavy loam, at elevations of 950 m.This site which lies east of the main winter rainfall region where Strumaria is centred, is located within a zone where the probability of rain is greatest in March (Zucchini & Adamson 1984).S. aestivalis responds rapidly to scattered summer thunder showers and flowers during January and February.
Stamens suberect to slightly spreading, exserted beyond the tepals; filaments separate, up to 17 mm long, with the outer and inner whorls adnate to the style base for up to 2.5 mm and 3.5 mm respectively; anthers subcentrifixed, ± 3 mm long before opening, dark maroon; pollen creamcoloured.Style up to 19 mm long, equalling or slightly exceeding the stamens, slightly thickened and trigonous proximally, tapering gradually upwards; with nectar col lecting in 3 droplets between the style and inner filaments; stigma shortly trifid.Seeds fleshy, ovoid, 2.0-2.5 mm diam., green to reddish brown.Chromosome number: 2n = 20.
Flowering time: May.but commencing in April when cultivated.
Diagnostic features: the long, lorate.pubescent leaves and somewhat funnel-shaped flowers of S. perryae are characteristics also found in S. pubescens and S. aestivalis, and indicate a close affinity with these species.The narrow leaves of S. perryae are diagnostic (at most 5 mm across).In contrast.5. pubescens and S. aesti\<alis have leaves more than 10 mm wide and the synapomorphy of yellow inner bulb tunics.The adnation of the filaments to the style is well developed and reaches a length of 3.5 mm.This feature is also conspicuous in specimens of S. pubescens, S. watermeyeri L. Bolus, as well as S. aestivalis.Unlike 5. aestivalis the inner filaments of these species are closely adnate to the style and the three efferent canals, which conduct nectar from the septal nectary to the sinus between the inner filaments and style, are only microscopically visible.
Distribution and habitat: S. perryae is known from a single small population on the northern Bokkeveld escarpment between Grasberg and Theunisdrift, north west of Nieuwoudtville (Figure 2).Plants grow in clay soil in association with low karroid shrubs.

Flowering time: March to May.
Diagnostic features: in comparison to the group of closely allied pubescent-leaved species with white-fleshed bulbs and stellate flowers, S. discifera has consistently long, narrowly lanceolate leaves, distinctly channelled tepals and a conspicuous bulbiform to discoid swelling at the base of the style.
Distribution and variation: Strumaria discifera is distributed between Vanrhynsdorp and Nieuwoudtville eastwards to Calvinia and the Roggeveld escarpment in the northwestern Cape (Figure 5).
The species includes a polymorphic range of popula tions.From the dolerite ridges on the outskirts of Nieuwoudtville the bulbs are densely clump-forming, whereas other known populations comprise scattered soli tary bulbs.The shape of the swelling at the base of the style is also variable.The clump-forming bulbs have a pronounced discoid stylar swelling with a frilly rim.This character state is fairly consistent within the population and is probably maintained through recurrent vegetative propagation.Collections east of Nieuwoudtville to the Hantamsberg and Bloukranz Pass near Calvinia also have disc-like swellings, but these are not as broad as those in the Nieuwoudtville populations and lack a prominent rim.Elsewhere in the distribution range the stylar swelling tends to be bulbiform in shape.Since the specimens from the clonal population on the dolerite koppies at Nieuwoudt ville can be adequately diagnosed, these are described here as a new subspecies.3a. S. discifera subsp.discifera.Figure 4.
Distribution and habitat: the known distribution extends from near Vanrhynsdorp, eastwards onto the Bokkeveld escarpment, across the high-lying plateau to Calvinia.then southwards along the edge of the Roggeveld escarpment to near Middelpos (Figure 5).Occupying gentle slopes and depressions, the taxon inhabits heavy loamy soils, most commonly derived from Nama and Ecca shales.The bulbs often grow in association with renosterbos (Elytropappus rhinocerotis (L.f.) Less.).with a prominent irregular rim on the disc distally, abruptly narrowed into a slender column above.
Distribution and habitat: subsp.bulbifera inhabits slopes and hollows of low exposed dolerite ridges on the Bokke-
Flowers 8-14, spreading, stellate, pure white or white to pale pink with a pale pink median dorsal stripe on each tepal, scentless; pedicels straight to upwardly curved.3 5 -80 mm long, concolorous with the scape.Tepals free to the base, outspread to slightly deflexed, oblonglanceolate, 6.5-8.5 x 2 -3 mm, distinctly channelled with slightly undulate margins proximally.Stamens equalling or slightly shorter than the tepals, spreading; filaments separate, adnate to the broadened style base, with the inner whorl attached higher up than the outer, broad but not bulbous basally, tapering slightly upwards; anthers subcentrifixed, 1.5 mm long and wine-red before dehiscing; pollen whitish.Ovary with 1-4 ovules per locule.Style up to 5 mm long, more or less equalling the stamens.
Flowering period extends from March to April.
Diagnostic features: the leaves of S. villosa are softly villous on the adaxial surface and are characteristically glaucous.Unlike other pubescent-leaved species of the Strumariinae with white, stellate flowers and filaments adnate to the style.S. villosa is specialized in having yellow inner bulb tunics.
Distribution and habitat: this rare species is known from only one locality in the Richtersveld.near Kosies, (Figure 5).Locally abundant on low hills, the species is confined to exposed, east-facing slopes amongst quartz pebbles which overlie weathered granite soil.initially green, breaking off at the base in fruit; spathe valves linear-lanceolate, 2 0-40 x 3 -7 mm; bracteoles filiform, up to 25 mm long.Flowers (2 0 -) 30-65, spreading, stellate, white to delicate pink with deep pink or greenish median stripes on the undersurface, ageing to light brown, with a heavy coconut-like scent; pedicels straight, 2 0 -5 0 mm long, becoming straw-coloured.Tepals almost free to the base or very shortly adnate to the staminal tube for up to 0.25 mm, otherwise outspread, oblong-lanceolate, 8-15 x 2 -4 mm. with plane edges.
Stamens equalling or up to 2 mm longer than the tepals, becoming outspread; filaments connate proximally into a tube protruding from the perigone throat by (1.0-) 1.5-4.0mm, subulate above, occasionally shortly toothed in the axils between adjacent filaments; anthers centrifixed, 3 mm long and dark w ine-red before opening: pollen creamcoloured.Style up to 15 mm long, narrow throughout, with nectar collecting in a well around the base; stigma shortly trifid.Seeds not known.Chromosome number: 2n = 22.cence; and the very short perigone tube (0.25 mm or less).
In these respects H. speciosa is similar to the shortly pubescent-leaved species, H. pilosula D. & U. Miiller-Doblies and H. incana Snijman, with which it also shares plane tepals.However, the glabrous leaves and the rela tive length of the stamens to the tepals distinguish it from these species.The stamens equal to or up to 2 mm longer than the tepals in H. speciosa, whereas they are distinctly shorter than the tepals (by 3 mm or more) in H. pilosula and H. incana.The inflorescence of H. speciosa may sometimes be confused with flowering material of H. breviflora Herb, from Namaqualand.but unlike this species the bulbs are without a conspicuous, exserted, red cataphyll which sheathes the foliage leaves.
Distribution and habitat: Hessea speciosa is recorded from red sand dunes and flats of friable loam, associated with the extensive drainage system of seasonal rivers from Warmbad in southern Namibia to Fraserburg in the central Cape.The associated vegetation is predominantly grassveld (Figure 9).
Variation: often the northerly populations have a distinct staminal tube (1.5-3.5 mm long), whereas specimens from the south of the distribution range have only a shortly developed staminal tube (less than 1.5 mm).Both white and pale pink flower forms occur, as well as the occa sional novelty of small teeth in the axils between adjoining filaments.
lanceolate, 3 0 -5 0 X 5 -7 mm; bracteoles filiform, up to 20 mm long.Flowers 10-20, spreading, widely funnelshaped, white, with a pale pink median dorsal band on each tepal, turning deeper pink with age, heavily scent ed; pedicels straight to upwardly curved, 40-55 mm long, pale greenish-pink.Tepals free to the base, spreading, oblong-lanceolate, 12-14 x 3 -5 mm.Stamens equalling the tepals, spreading slightly from near the base; filaments separate, 7-10 mm long, adnate to the broadened style base for up to 4 mm; the inner face of the inner whorl free with only