Albuca crispa and A. grandis (Hyacinthaceae: Omithogaloideae), two new species of subgenus Albuca, the rediscovery of A. albucoides (sub genus Osmyne), and the identity of A. reflexa

Albuca crispa is a new species o f section Falconera series Trianthera w ith crispulate leaves from the Great Karoo, know n at least since 1947 but overlooked until now. A second new species. A grandis. from the southwestern Cape was prev iously included in A. fragrans Jacq. (section Falconera series Falconera). It is a robust species that flowers in w inter and early spring and the styles are rugulose w ith ± isodiametric epidermal cells, unlike typical A. fragrans w hich is a more slender species flowering in early summer and w ith derived, smooth styles w ith fusiform epidermal cells. The recent discovery o f a flowering population matching the type o f A. albucoides (Aiton) J.C.Manning & Goldblatt (subgenus Osmyne) allows for a full description and illustration o f this poorly know n and taxonomically neglected species that has often been included in A. suaveolens (Jacq.) J.C.Manning & Goldblatt. Lastly, examination o f the type o f A. reflexa Krause & Dinter from Namibia shows it to be conspecific w ith Drimia indica (Roxb.) Jessop.


INTRODUCTION
Hyacinthaceae, which comprises 700-900 species o f bulbous geophytes widely distributed through Africa and the Mediterranean, extending to Northern Europe.Asia and South America, is one o f the larger geophyte fami lies in southern Africa.The division o f the family into the four subfamilies Hyacinthoideae.Omithogaloideae.Oziroëoideae and Urgineoideae (Speta 1998: Manning el al. 2004) is now well established but generic circum scriptions in subfamily Omithogaloideae in particular have undergone major revision since 1998 (Speta 1998: Manning et al. 2004Manning et al. , 2009)).The description o f Orni thogalum L. (Linnaeus 1753) was followed by that o f Albuca L. (Linnaeus 1762), which was established to accommodate those species o f Ornithogalum w ith dis tinctive Galanthus-Uke flowers in which the outer tepals are ± spreading and the inner erect and ± connivent.This distinction has been readily applied in most instances, with the notable exception o f members o f Ornithogalum subgen.Osmyne (Salisb.)Baker (Obermeyer 1978).The flowers o f members o f this group have fleshy, yellow ish tepals with a dark median band that are highly remi niscent o f flowers o f Albuca.leading early workers to describe several species o f this taxon in the latter genus.
The recent application o f DNA techniques to the fam ily suggested that neither Albuca nor Ornithogalum w ere monophyletic as then defined, leading Manning et al. (2004) to include both genera within a greatly expanded Ornithogalum.further analysis provided greater reso lution o f relationships within the subfamily, enabling Manning et al. (2009) to develop a revised classification of the subfamily in which the genus Albuca is reinstated in a modified circumscription that includes several ele ments previously assigned to Ornithogalum.notably subgen.Osmyne and most members o f subgen.Urophyllon (Salisb.)Baker.In this circumscription Albuca is distinguished from Ornithogalum by thick-textured tepals with a broad median green band associated with a concentration o f vascular traces around the midline, the style mostly longer than the ovary, and the generally larger seeds.This classification is followed here.

Subgenus Albuca
Species o f Albuca subgen.Albuca are distributed among the four sections Albuca.Falconera (Salisb.)J.C.Manning & Goldblatt.Branciona (Salisb.)J.C.Manning & Gold blatt (= sect.Mitrotepalum U.Mull.-Doblies ms.) and Pallastema (Salisb.)J.C. Manning & Goldblatt (Manning et al. 2009).The circumscription of these taxa at the level of subgenera was largely established by Muller-Doblies (1995).whose partial accounts o f the species (Muller-Doblies 1994. 1995. 2006) constitute the most recent revi sions of the group.Falconera, in contrast to that in other sections in the subgenus, is relatively diverse, which enabled Muller-Doblies (1995) to identify significant interspecific varia tion in androecial and especially gynoecial morphology.Vegetatively the species in the section are almost evenly divided between those with leaves (and often peduncle) variously covered with glandular trichomes, and those with the vegetative parts completely glabrous.Among the latter, Albuca fragrans Jacq.and A, clanwilliamaegloria U.Mull.-Doblies are considered to be sister spe cies on the basis o f their unique stylar morphology.In most species o f subgen.Albuca the epidermal cells of the style are ± isodiametric but in these two species they are fusiform, giving the style a distinctive, longitudinally striate surface under magnification.This condition was first identified by Miiller-Doblies (1995), who described the cells as prosenchymatous, a term originally applied to tissues o f elongate cells concerned with the special ized functions o f support, protection or conduction, in contrast to parenchyma, which is concerned with more generalized vegetative aspects.The term is thus conven tionally applied to the vascular tissue and to supporting sclerenchymatous strands and although now generally obsolete, it is still used to described cells that are elon gate and taper-pointed, thus spindle-shaped or fusiform (Eames & MacDaniels 1925).
Albuca clanwilliamaegloria is one o f the most distinc tive species in the genus (M anning et al. 1999), readily recognized by its height and large, dull yellow flowers w ith suberect tepals.A. fragrans, in contrast, is far less distinguished and has until now included two very dif ferent elements.The first comprises slender, late-flower ing plants with the distinctive fusiform stylar cells that characterize the species.The second element consists o f robust plants that flower in w inter and early spring and have a rugulose style with ± isodiametric epider mal cells.The latter plants are common and conspicuous along the West Coast and have been illustrated under the name A. fragrans in several popular accounts o f the flora (Manning & Goldblatt 1994;Manning et al. 2002).
Distribution and ecology-: Albuca crispa appears to be endemic to the Great Karoo, where it has been found between Beaufort West, Graaff-Reinet and Steytlerville (Figure 2).It has been recorded on rocky flats, and flow ers in midsummer.

Distribution and ecology': distributed along the West
Coast o f Western and Northern Cape, with collections from north o f Hondeklip Bay in central Namqualand as far south as Koeberg (Figure 2).The species is poorly collected despite, or more probably because of, its large size, and the exact extent o f its range has still to be deter mined.Albuca grandis occurs in deep sandy soils, usu ally in coastal thicket and scrub where plants are often sheltered by bushes and shrubs.
Diagnosis and relationships: Albuca grandis is a robust species, typically reaching a metre or more in height, with a characteristic thick peduncle, usually more than 10 mm in diameter, and a dense, conical raceme of large, dull greenish white flowers with tepals 22-27 mm long.All six stamens are fertile, although the anthers o f the outer whorl are ± one third smaller than those o f the inner whorl.The ovary is conspicuously 3-lobed, almost winged, with parallel paraseptal ridges, and a rugulose style.The bulb tunics are not at all fibrous.A. grandis is distinguished from other glabrous species o f series Fal conera with rugulose styles by its stature and the rela tively early flowering, mainly in August and September.
The species has until now been confused with Albuca fragrans but the two differ in details o f morphology and in ecology and phenology.Generally less than 800 mm tall, A. fragrans may reach up to I m but is always a slender plant with the rachis at most 5 mm in diameter.A lbuca fra g ra n s Jacq., Plantarum rariorum horti caesarei schoenbrunnensis 1: 44 (1797).Type: South Africa, [Western Cape], without locality or date, illustra tion in Jacquin, Plantarum rariorum horti caesarei schoen brunnensis 1: t. 84 (1797).
Stamens adnate to perianth for ± 1 mm.dimorphic, erect; outer filaments narrowly lanceolate-subulate, 12-14 x 2 mm with anthers ± 4 mm long at anthesis, cream-coloured.inner filaments laterally expanded and pinched in lower fourth, ±12x2 mm with anthers ± 6 mm long at anthesis, cream-coloured.Ovary' weakly hourglass shaped, 3-lobed, green, 6-8 mm long, shortly stipitate, paraseptal ridges parallel; style columnar, trigonous, longitudinally striate with testal cells fusiform, yellow ish green, 8-9 mm long, obtuse with trigonous, papillate    several, mostly narrow and not highly succulent leaves.Distinctions between many o f these species are based primarily on differences in leaf number, orientation and morphology.The taxonom ic treatment o f the group by Obermeyer (1978) was hampered by inadequate collec tions o f many o f the taxa, with the result that species were often treated rather broadly.A more recent tax onomy o f the group (M uller-Doblies & Miiller-Doblies 1996) has benefited from much more extensive field knowledge but many o f the species are not supported by adequate descriptions nor, in several cases, by the depo sition o f type specimens, making their assessm ent diffi cult if not impossible.
An extensive population o f autum n-and early winter-flow ering plants from the low lands at the foot of the Elandsberg M ountains between M almesbury and Tulbagh in W estern Cape could not be assigned to any o f the species o f the subgenus generally recog nized from the West Coast, and it was initially con sidered that they m ight represent an undescribed spe cies.The leaves o f these plants, which are absent or incom pletely developed at flowering, num ber between 3 -5 , are erect and ± subterete or terete, and the abaxial surface is adorned with longitudinal row s of minute papillae.Sem i-terete form s have a translucent adaxial band down the m iddle o f the blade and the margins are scabridulous.The flowers are suberect but otherwise unexceptional in the group.
More than half o f the species in subgen.Osmyne are confined to Namaqualand, with just five o f the currently named species recorded in the southwestern Cape south o f Clanwilliam.Among these, only Albuca albucoides (Aiton) J.C.M anning & Goldblatt (= Ornithogalum albu coides (Aiton) Thunb.)approaches the Elandsberg plants in having subterete or semiterete leaves, and is also reported to be hysteranthous.Non-flowering plants that are attributed to the species have been widely recorded by Muller-Doblies & Muller-Doblies (1996) along the West Coast, from the Bokkeveld Mountains to the Cape Peninsula but no specimens have been located in South African herbaria.Known in flower solely from the type, the species was included in A. suaveolens (Jacq.)J.C.M anning & Goldblatt by Obermeyer (1978)  A lbuca albucoides (Aiton) J.C. Manning & Gold blatt in Taxon 58: 94 (2009).Anthericum albucoides Aiton: 449 (1789).Ornithogalum albucoides (Aiton) Thunb.: 62 (1794).Type: South Africa, [Western Cape], without precise locality or date, Masson s.n.ex cult, hort.Kew 1788 (BM.holo.!).
Albuca sect.Falconera comprises 18 species divided between the two series: Trianthera (U.Mull.-Doblies)J.C.Manning & Goldblatt (the outer whorl o f stamens sterile): and Falconera (Salisb.)J.C.Manning & Gold blatt (both staminal whorls fertile) (Muller-Doblies 1995: Manning & Goldblatt 2006: Manning et al. 2009).Series Trianthera comprises just five species from eastern and southwestern South Africa.Several collections o f a dis tinctive member o f this series from the Great Karoo w ith unique, crispulate leaves represent an unnamed species that we describe here as A. crispa.making it the sixth member o f the series.Series Falconera.the largest o f the two series in subgen.Falconera.currently includes 13 species from southern and south tropical Africa but is largely confined to the winter rainfall region o f southern Africa, includ ing southwestern Namibia.Floral morphology in sect.
Among the remaining species o f sect.Falconera w ith glabrous leaves and rugulose styles, Albuca papyracea J.C.M anning & Goldblatt and A. robertsoniana U.Mull.-Doblies are characterized by a papery or fibrous neck sheathing the base o f the stem and both are currently known only from the Robertson and Little Karoos; A. bifoliata R.A.Dyer from Grahamstown has unique, recurved stigmatic horns; A. monophvlla Baker from southern Angola and Namibia has a single leaf and a style that is at least tw ice as long as the ovary; and A. kirstenii (J.C.Manning & Goldblatt) J.C.Manning & Gold TYPE.-Western Cape, 3222 (Beaufort West): Karoo National Park Camp at main entrance gate, 840 m, ( -BC), 9 December 2005, Bester 6220 (NBG.holo.; PRE.
but was resurrected by MQller-Doblies & Muller-Doblies (1996).In the absence o f any further collections, however, we continued to include the species in A. suaveolens (M an ning & Goldblatt 2006).Albuca albucoides is based on a single plant collected by Francis Masson from the Western Cape, most likely from either the Paardeberg near Malmesbury or from near Piketberg according to the conclusions o f Muller-Doblies & Muller-Doblies(1996).The plant flowered in cultivation in Britain and the type specimen comprises five detached leaves and a single raceme.The date of flowering is given as September and October.This is almost certainly based on its flowering in cultivation in the northern hemisphere, which would correlate to March or April in the southern hemisphere.The leaves o f the species are very characteristic at close inspec tion (Muller-Doblies & Muller-Doblies 1996), being semi-terete, 2 -6 mm wide, stiff and succulent, shallowly canaliculate above, with an asperulous margin and with papillate ridges on the dorsal (sic) surface.This descrip tion is a perfect match for the Elandsberg plants, apart from the location o f the papillate ridges on the morpho logically ventral surface.Examination o f the type speci men confirms that it matches the Elandsberg plants very well in the leaves and also in the few-flowered raceme with suberect flowers.There is thus no doubt that the Elandsberg plants represent the poorly understood Albuca albucoides and we take this opportunity o f pro viding a full description and illustration o f this Most' spe cies.
Distribution and ecology", thus far collections o f flow ering material o f Albuca albucoides have been made at only two sites, one the type made in the late eighteenth century and the other the recent collections from Elands berg Farm at the western foot o f the Elandsberg Moun tains east o f Riebeeck-Kasteel (Figure 6).The type is thought to have been collected either on the Paardeberg near Malmesbury or near Piketberg (Muller-Doblies & Muller-Doblies 1996), and the two fully documented localities for the species are thus both from the small area along the West Coast known as the Swartland, between Malmesbury and Piketberg.Nonflowering plants, which were assigned to the species, have been collected from several additional localities along the West Coast, ranging from near Mamre in the south to Nieuwoudtville in the north (Muller-Doblies & Muller-Doblies 1996).The plants at Elandsberg occur in local colonies on stony alluvium in the highly localized Swart land Alluvial Fynbos vegetation (Mucina & Rutherford 2006).The nonflowering plants assigned to the species have been collected primarily from rock ledges in fyn bos but also from deep sands and gravelly flats.The full extent ol the range and ecology o f A. albucoides remains to be adequately documented.Flowering in the Elandsberg population takes place in the late autumn and early winter, from late March to early May, and we have observed more than one flush o f emer gence, presumably under the influence o f environmental stimuli such as the passage o f cold fronts.Early flower ing plants appear without the leaves but those that are stimulated into flower later in the season are accompa nied by the appearance o f a tuft of young leaves lateral to the flowering stem.Seeds are shed soon after flowering, within two or three weeks, and thus plants with dehisced