A revision of Ornithogalum subgenus Aspasia section Aspasia ^ the chincherinchees ( Hyacinthaceae )

The species o f Ornithogalum L. subgenus Aspasia section Aspasia are revised. Section Aspasia is defined by a rosette o f lanceolate to oblong leaves; large, boat-shaped, ± petaloid bracts; moderately-sized white, yellow or orange flowers, sometimes with dark central markings; thin-textured, ellipsoid capsules that are enclosed by and concealed within the persistent, papery perianth; and angular, colliculate to echinulate seeds. Twelve species are recognized in the section, separable into three series based on seed morphology. O. conicum is redefined to exclude specimens from the Eastern Cape, which are recognized as O. synanthifolium, and O. conicum subsp. strictum is raised to species status as O. strictum. The circumscription o f O. duhium is expanded to include O.fimbrimarginatum and O. subcoriaceum, previously distinguished on account o f their longer styles. Collections from the Roggeveld Escarpment and Klein Roggeveld that were previously included in O. fimbrimaiginatum are recognized as the new species O. corticatum Mart.-Azarin, on the basis o f their unusual, thick, cartilaginous outer tunics and puberulous adaxial leaf surface. O. ceresianum is removed from the synonomy o f O. thyrsoides and recognized as a distinct species on account o f its extensive glossy black tepal markings, winged inner filaments, and glossy black ovary. The poorly known O. puberuhim is more ftilly described based on several recent collections, and O. leeupoortense is neotypified in the absence o f any original type material. O. rupestre and O. multifolium are regarded as colour forms o f the same species, for which O. rupestre is the older name. Similarly, O. roussouwii is a depauperate, pale form o f O. maculatum and is thus included in the synonomy o f that species. The circumscription o f O. pruinosum remains unchanged. The species O. haurii, O. diphyllum and O. sephtonii from the Drakensberg Mountains o f Eastern Cape and KwaZulu-Natal are excluded from section Aspasia on the basis o f their turbinate capsules that are exposed by the reflexed tepals. Each species is fully described, with accompanying discussion o f variation and relationships, a distribution map, and an illustration. Bothalia 37,2: 133-164(2007)


INTRODUCTION
Hyacinthaceae, a predominantly Old World family, is distributed throughout Africa and the Mediterranean, extending through central to eastern Asia, with a single, small genus in the Andes Mountains in South America (Speta 1998).Around 400 of an estimated total of 700-900 species are endemic to southern Africa, making it one of the most important geophyte families in south ern Africa, exceeded in numbers only by the Iridaceae.Hyacinthaceae are especially numerous in the south western winter rainfall region, which is one of the main centres of diversity for the family.
Among the more common and conspicuous members here are the large-flowered species of Ornithoglaum L. subgenus Aspasia (Salisb.)Obenn., known colloquially as chincherinchees, an onomatopoeic sobriquet derived from the sound produced when the stems are rubbed together (Smith 1966).Several species in this group, especially O. thyrsoides, are a characteristic part of the southwestern Cape spring, occurring in enonnous popu lations in lowland areas around Cape Town and further north into Namaqualand.Their extreme toxicity (Van Wyk et al. 2002) enables them to colonize overgrazed lands with impunity, and their ease of cultivation and long vase-life have made them important horticulturally.Several selections of O. dubium and O. thyrsoides are available commercially as cut-flowers and further breeding programmes are ongoing.The taxonomy of this group, however, is far from fully understood and several of the species are notoriously difficult to identify.
The southern African species of Ornithogalum have been revised three times in the last sixty years, beginning with the work of Leighton (1944Leighton ( , 1945)).This review was the first complete regional treatment o f the genus since Baker's (1897) account for the Flora capensis, with the apparently significant advantage gained from examina tion of living material collected in the wild.Leighton, like Baker before her, was impressed by the great vari ability among populations in the form of the inner fila ments, which may be slender and awl-like, lanceolate, with or without small basal expansions, or conspicuously winged.Flower colour is another source of variation, ranging from pure white through various milky-white or buff shades to yellow, orange or reddish orange, with or without small to large dark central markings.Defining her taxa very narrowly, Leighton (1944Leighton ( , 1945) ) described numerous species to accommodate the different combi nations of these characteristics, ultimately recognizing 21 species and many forms in the group (Leighton 1945).With further study, however, it became clear that many of these species represented nothing more than local populations.Obermeyer (1978), who was the first to pro pose a more formal recognition of the chincherinchees, as the Aspasiae group of subgenus Aspasia, adopted a much broader view of the species in her monograph on the genus, reducing their number to 10, with two subspe cies recognized in O. conicum.Ornithogalum diphyllum Baker from the KwaZulu-Natal Drakensberg, unaccount ably left out of Leighton's treatment, was included as the eleventh member of the group.This treatment was largely followed by Miiller-Doblies & Miiller-Doblies (1996), who formalized the group as section Aspasia.The sec tion was expanded slightly to include two additional species, O. constrictiim F.M.Leight.and O. inclusum F.M.Leight., which had been placed by Obermeyer in the Hispidaspasiae group, and was subdivided into two subsections and six series.The species in Obermeyer's Aspasiae group were dispersed among four series in two subsections: Aspasia, Leeupoortensia U. Despite the relatively high level of taxonomic study to which it has been exposed, the taxonomy of the group remains unclear, and the identification of specimens is often problematical.Although some of the species are well circumscribed and may be identified without much difficulty, the boundaries of others, especially those around O. dubium, remain poorly defined.Increased col lecting over the past decades has greatly improved our appreciation of the natural variation among wild popula tions, indicating the need for another review of the spe cies in this group.A comprehensive, illustrated account of the species is presented here for the first time.
It is evident that many of the characters that have tra ditionally been used to separate species in the group are much more variable than has been realized.This is par ticularly true of flower colour, the degree of basal expan sion of the inner filaments, the length of the style rela tive to the ovary, and the surface sculpturing of the seeds.Most species are reliably diagnosed by a combination of characters, supplemented with distribution and ecological data.A previously underappreciated character, the size of the seeds, appears to be a useftil indicator of relation ships among the species.Three seed size classes can be distinguished: large (2.0-3.5 mm long), in O. conicum, O. corticatum and O. synanthifolium\ medium (1-2 mm long), in O. ceresianum, O. strictum and O. thyrsoides; and small (0.5-1.0 mm long), in the remaining species.The species are arranged in three series according to these size classes.

MATERIALS AND METHODS
This study is based on an examination of dried her barium specimens as well as living plants studied during extensive field work undertaken thoughout the southwestem Cape.The herbarium specimens studied include the complete collections in BOL, K, NBG and SAM, the types of all names, and selected specimens from PRE. Specimens examined are listed at the end.Seeds were examined with both light and scanning electron microscopy.TAXONOMY Ornithogalum L., Species plantarum: 306 (1753).Type : O. umbellatum L. (vide Steam 1983).
Deciduous or rarely evergreen perennials.Bulb subter ranean or epigeal, subglobose or rarely poorly developed and rootstock rhizomatous, tunics sometimes scale-like or loosely overlapping, usually white but rarely pinkish; outer bulb tunics membranous, papery, or leathery.Leaves 1-several, green or dry at flowering, erect or spreading, linear to oblong or filiform, sometimes very succulent, usually glabrous but sometimes pubescent or glandularpubescent, margins smooth, ciliate, fringed, or hyaline, sheaths sometimes persistent and forming a papery or weakly to strongly fibrous, sometimes horizontally barred sheath around base of stem.Inflorescence a several-to many-flowered raceme, sometimes subcorymbose with shortened axis, or secund, usually solitary but sometimes more than one; peduncle rarely papillate; bracts membra nous, leafy or petaloid, small or large, not spurred; bracteoles usually lacking but sometimes thread-like and bome on alternate sides of pedicels at base; pedicels short or long.Flowers white, yellow, orange, or yellowish green, without darker keels, scented or unscented, sometimes closing at night or more rarely nocturnal, suberect or pat ent, rotate or campanulate; tepals ovate to narrowly lan ceolate, persistent, ± shortly united at base or rarely united into a short tube, spreading to erect, sometimes with tips reflexed.Stamens suberect or slightly spreading; fila ments free or rarely united below, fused to base of tepals or inserted at top of tube, filiform to lanceolate, all similar or inner usually broader, both whorls or more usually only inner variously expanded or toothed below.Ovary ovoid to globose or turbinate, sometimes shortly stipitate; ovules few to many per locule; style vestigial to long, subcylindrical, erect or sometimes slightly deflexed; stigma small and 3-lobed or -angled.Capsule fiisiform, ovoid to subglobose, ± 3-angled or -lobed, membranous or leathery, enclosed in dry perianth or exposed, dehiscing loculicidally.Seeds few to many per chamber, flattened or angled, black, the testa tightly adhering, laevigate, rugulate, papillate or echinate.Base chromosome number x = 9.About 250 species in Africa, Madagascar, Mediterra nean, Saudi Arabia and India, mainly in the winter rainfall areas of southern Africa, including ± 40 species in Western Cape, South Africa.
Subgenus Aspasia (Salisb.)Oberm. in Bothalia 12: free, ovate, persistent and papery in fruit.Stamens free;333 (1978) ,hin-walled, enclosed and concealed within dry but sometimes partially or entirely pubescent, mar-perianth.Seeds many per chamber, pyriform, commagins usually fringed or ciliate.Inflorescence sometimes shaped or cuneate, 1-3 mm long, testa papillate or echisubcorymbose; bracts large, boat-shaped, foliaceous nuU e. Base chromosome number x = 6.or petaloid, margins entire or ciliate but not denticu late.Flowers medium-sized to large, white, yellow or Species 12, mainly endemic to the winter rainorange, without darker keels but sometimes with dark fall region of the southwestern Cape, with tw^o species centre, closing at night, shallowly bowl-shaped; tepals extending eastwards to the Eastern Cape.

owt ■
Distribution and ecology: restricted to coastal areas in the southwestern Cape, from Graafwater southwards along the west coast to the Cape Peninsula and Gordon's Bay (Figure 3), occurring on coastal limestone pavement, stony hills and granitic outcrops, typically in coarse grained sandy or gravelly soils but also shale.
Discussion: Omithogalum conicum is a summer-flow ering species from coastal areas in the extreme southwest ern Cape.It is characterized by its very large bulb, some times with pale pink flesh, whitish to light grey, papery tunics, and a rosette of short, densely ciliate leaves less than one third as long as the inflorescence and partially or completely dry at flowering.Both whorls of filaments are typically filiform or awl-shaped, or the inner may rarely be slightly widened at the base but never into the apically lobed, oblong expansions that occur in other species in the group.which often forms large colonies along road verges and in waste ground, differs in its much smaller bulb, seldom more than 20 mm in diameter, longer, suberect, ± synanthous leaves, and conspicuously winged inner filaments.It also flowers earlier in the season, in September and October, and is in fruit by the time that O. conicum begins to bloom in early November.White-flowered forms of O. dubium that occur on Table Mountain are distinguished by their smaller bulb with dark tunics, mostly subcorymbose inflorescence, and basally expanded or winged inner filaments.In addi tion, the ovary in O. dubium is typically blackish, unlike the yellow ovary in O. conicum, and the seeds are much smaller (± 1 mm vs 2-3 mm long).On the Peninsula, O. dubium is restricted to sheltered places on sandstone cliffs, whereas O. conicum is found at lower altitudes on exposed granite or shale slopes.The flowers in the populations of O. conicum on Lions Head on the Cape Peninsula are open throughout the day from early morn ing, and are slightly scented, unlike those of O. dubium, which only open around mid-morning and lack fra grance.
Although the species has been documented as toxic, the actual identity of the specimens tested is uncertain in view of the much broader circumscription of the species that was current in the past.Given its relatively restricted geographic range, however, it seems more likely that these results were based on tests of O. strictum rather than true O. conicum.History'-.Ornithogalum conicum was illustrated and described by Jacquin fi-om cultivated plants that were almost certainly originally collected on the Cape Penin sula, where the species is still found on the slopes of Signal Hill and the foot of Lions Head.At the same time, Jacquin illustrated and named O.lacteum but this appears to be nothing more than a luxuriant form with the bases of the inner filaments slightly expanded.This form was reduced to synonomy by Baker (1897), whose concept of O. conicum, later followed by Obermeyer (1978) Plants (200-)350-950 mm high.Bulb ovoid, some times not well developed, (20-)25-35 mm diam., outer tunics not always persisting but then greyish to blackish, leathery.Leaves suberect, 5-11, synanthous, oblong-lan ceolate, ± half as long as inflorescence, (150-)200-400 X (8-)10-25(-30) mm, dark green, soft-textured, gla brous or ciliolate on margins.Raceme subcorymbose or cylindrical, compact, (50-)70-150 mm long but elon gating to 100-150 mm long in fruit, (10-)15^0(-50)flowered, sometimes two per bulb; lowermost pedi cels 15-30 mm long, elongating to 35-72 mm in fruit; bracts pale and papery, ovate, acuminate, lowermost (12-) 15-40 mm long, longer or shorter than lower pedi cels in flower and shorter in fruit.Flowers white; outer tepals ovate-lanceolate, inner obovate-lanceolate, 13-18(-20) X 6-8(-10) mm.Stamens half as long as tepals; filaments 5-7 mm long, white, outer linear to narrowly subulate, inner either subulate or expanded and oblong in lower half, rarely winged.Ovary' ovoid, 3-5 mm long, greenish; style erect, 2-3 mm long.Capsule oblongovoid to obovoid, 3-lobed, obtuse-apiculate, 12-15 mm long.Seeds angular, irregularly folded, 2.5-3.5 mm long, colliculate to papillate, especially along margins.Chromosomes: 2n = 12 (De Wet 1957;Pienaar 1963).Flowering time: September to January.Figures 1B, 4.  Distribution and ecology: endemic to the Eastern Cape, where it is scattered along the higher ground fringing the coast, from Umtata in the north to Grahamstown in the south (Figure 3), occurring in moist or marshy grassland along streams and in damp valleys, sometimes in dense colonies.The species is typically evergreen and flowers throughout the wet summer season.
Discussion: Ornithogalum synanthifolium is typically a large, often evergreen species 350-950 mm high with relatively long leaves, usually at least half as long as the inflorescence.The leaves are soft-textured and bright green, with the margins glabrous or at most ciliolate.The plants produce a subcorymbose to cylindrical raceme of pure white flowers with elongate lower pedicels, 15-30 mm long in flower but lengthening to 35-72 mm in fruit.It is not uncommon for the plants to produce two inflor escences in a single season (e.g.Pienaar 8) or for the inflorescence of the previous season to persist alongside that of the current year due to the ± continuous vegeta tive growth in the species.
The bulb in O. synanthifolium may not be well devel oped as a result of both its evergreen habit and its pref erence for moist habitats, and the outer bulb tunics may not persist for the same reasons.When present, the outer tunics are greyish or blackish and somewhat leathery in texture.The inner filaments, like those o f most species in the section, are rather variable in shape, ranging from lanceolate to basally expanded or winged, sometimes {Taylor 4212) to the degree that they resemble those of O. thyrsoides.The large capsules, 12-15 mm long and large seeds, 2-3 mm long, are shared with O. conicum.Other members in the section have smaller capsules, and seeds less than 2 mm long.
There is no doubt that the Eastern Cape plants are quite distinct from O. conicum, which occurs in sea sonally dry situations on the Cape Peninsula and adja cent parts of the southwestern Cape.The climate here is Mediterranean and summer-dry, unlike the summer rain fall climate in which O. synanthifolium occurs.True O. conicum invariably produces a well-developed, subglobose bulb and a rosette of short, spreading, densely ciliate leaves that are dry and withered at flowering, which takes place in early summer after the rainy season.
In the wild, Ornithogalum synanthifolium may be confused with O. dubium, the only other species of sec tion Aspasia to occur in the Eastern Cape, but the lat ter is deciduous, grows in more rocky situations, and produces a rosette of spreading leaves that are densely ciliate on the margins and usually shorter than half the length of the inflorescence.The raceme in O. dubium is also typically shorter and more densely corymbose than in O. synanthifolium, and the seeds are much smaller, ± 1 mm long.
History: Ornithogalum synanthifolium has been much misunderstood in the past.The first collection of the spe cies appears to have been made by the Rev. F. Baur near Baziya in Eastern Cape.It was identified as O. lacteum Jacq.(now regarded as a synonym of O. conicum Jacq.) by Baker (1897) on account of its awl-shaped inner fila ments.Another early collection made by Ernest Galpin, a local businessman with an intense interest in botany, probably around the turn of the century, formed the basis of O. synanthifolium, which was described by Leighton (1944) for several collections of plants from the Eastern Cape that have the inner filaments expanded at the base but which otherwise closely match those collected by Baur.The latter she retained in O. lacteum, albeit as a distinct form restricted to the Eastern Cape.
Distribution and ecology': endemic to the Roggeveld and Klein Roggeveld near Sutherland in Northern Cape (Figure 3), where it favours heavy clay soils derived from dolerite, growing in open renosterveld shrubland.On the Roggeveld Escarpment, the species may occur in dense colonies numbering thousands of plants that cover large swathes of country.This suggests that it may be toxic to stock.On the more rocky slopes of the Klein Roggeveld, plants tend to be more scattered, sometimes on sandstone pavement along drainage lines.
The old bulb tunics of Ornithogalum corticatum are exceptionally firm, accumulating around the bulb in thick, rigid layers.Cartilaginous or thickly mat ted tunics are a feature of other geophytes from the Roggeveld Escarpment, including Lachenalia comptonii (Hyacinthaceae) and Devia xeromorpha (Iridaceae).The region experiences extreme climatic conditions through the year, with bitterly cold winters, during which tem peratures fall below freezing, contrasting with hot sum mers when temperatures reach the high thirties, and the thickened covering of the underground parts may act as insulation.
Ornithogalum corticatum flowers in early sum mer, at which time the leaves are completely withered.Shrinkage o f the bulb at this time results in the flesh pulling away from the stiff outer tunics, which form loose, easily detachable layers.
Discussion: Ornithogalum corticatum is readily dis tinguished by its depressed-globose bulb surrounded by firm, cartilaginous tunics, and rosette of short, spreading leaves, less than one quarter the length of the inflores cence, that are shortly and densely hairy on the upper surface in the distal half, and thickly fringed on the mar gins.These are completely withered at flowering but the pubescence is still visible.The pure white flowers are borne in conical or narrowly cylindrical racemes on short pedicels up to 12 mm long.The relatively large, strongly echinate seeds are also distinctive.Additional support for its recognition as a distinct species comes from the cytological studies of Pienaar (1963) and Roos & Pienaar (1966), which identified these plants as a dis tinctive form of O. lacteum (a synonym of O. conicum) with the unusual chromosome number of 2n = 10, other members of the section typically having 2n = 12.
Distribution and ecology: common along the Bokkeveld Mountains around Nieuwoudtville and through the Olifants River Valley, with outlying populations at the foot of the Piketberg and in the Ceres Valley in the south, and scattered collections ft-om the Roggeveld and Klein Roggeveld in the east, fi-om 100-800 m above sea level (Figure 8).The species occurs mainly on fine-grained clay or loam, typically on moist slopes or in drainage lines or vleis, where it can occur in large concentrations.On the Klein Roggeveld the species grows in washes along sea sonal streams.
Ornithogalum strictum is the common white chincherinchee in the Olifants River Valley, sometimes forming large swathes in seasonal vleis on the valley bottom.
Discussion: Ornithogalum strictum is recognized by its thin-textured, pale grey or whitish bulb tunics, long, erect leaves that often reach to the base of the raceme and are green at flowering, and by the narrowly conical raceme with short lower pedicels, typically less than 15 mm long.The flowers are usually pure white, although populations near Citrusdal may include individuals with a brownish centre.The ovary is mostly yellow or yellow ish green and the style is typically deflexed in a highly characteristic manner unique to the species.The inner filaments are usually widened in the lower third into an oblong expansion, sometimes bilobed above but not dis tinctly winged.As in most species of section Aspasia, O. strictum shows some variation in the degree of expan sion of the inner filaments.Plants ft-om the Bokkeveld Escarpment have the inner filaments invariably expanded in the basal third but populations from fiirther south, in the Olifants River Valley and around the Piketberg, may have almost linear or subulate inner filaments.
Ornithogalum strictum resembles O. thyrsoides in its pale outer tunics, slender leaves clasping the base of the stem, and expanded inner filaments, and both seem to occupy similar, seasonally moist habitats.O. thyrsoi des can usually be distinguished by its subcorymbose to broadly cylindrical inflorescence with longer lower pedicels, especially in ftiiit, and inner filaments that are broadly expanded in the lower half (not third) and always conspicuously winged, with the wings curved over the ovary, which is typically dark greenish or blackish.(Leighton 1944;Obermeyer 1978), although it continued to be distinguished at an infraspecific level by its slender, narrowly conical raceme.The decision to include it in O. conicum was made on account of the relatively weakly expanded inner filaments, which are quite unlike the broadly winged fila ments that are diagnostic of O. thyrsoides.It is now clear that O. conicum, with its large bulb, short, densely ciliate leaves, and larger seeds, is quite a different species, and on vegetative and seed characters it appears that O. stric tum is actually more closely allied to O. thyrsoides.
Distribution and ecology: widespread through the southwestern Cape, from Bredasdorp in the south to Vanrhynsdorp on the West Coast, with outlying popula tions recorded from further north on the higher ground in central Namaqualand, between Caries and Steinkopf (Figure 10).The species is typically found at lower alti tudes and has not been recorded from the Bokkeveld, Cedarberg or Cold Bokkeveld Mountain ranges.We have not encountered it in the Warm Bokkeveld either, and the only known collection from Ceres {Walters 904) prob ably refers to the general region and not the town itself.The species is common on shale or loamy soils, espe cially in seasonally moist sites and ditches along road sides.It appears to flourish under slight disturbance or overgrazing, and in such conditions may occur in dense colonies numbering hundreds or thousands of plants.
Ornithogalum thyrsoides is the most commonly encountered species in the southwestern Cape, especially along the west coast between Piketberg and Gordon's Bay, and is the original florists' chincherinchee.
It is known to be highly toxic to stock (Van Wyk et al. 2002), which may contribute to its rather weedy propen sity.The toxic compound in the plants has been identi fied as prasinode G, a steroid glycoside (Van Wyk et al. 2002).
Discussion: Ornithogalum thyrsoides is characterized by its relatively small bulb with pale, membranous outer tunics, and slender, erect leaves that clasp the base o f the stem.The outer filaments are slender and awl-shaped but the inner filaments are conspicuously expanded in the lower two-thirds, the expansions with prominent apical wings that are curved over the dark greyish to brownish ovary.O. thyrsoides is usually readily distinguished from other species, including O. conicum, by the combination of thin-textured, pale tunics and winged filaments but may be difficult to differentiate from O. strictum in the Olifants River Valley where the two species co-occur.O. strictum typically has a longer, narrower raceme, inner filaments that are only expanded in the lower third and not winged, and an ovary that is usually yellow or pale greenish with a deflexed style.
Further south, in the Swartland and Boland, Ornitho galum thyrsoides occurs with O. dubium but that species is distinguished by its black outer tunics, shorter, more densely cilate leaves and often shorter style.In addition, the filaments in O. dubium are often thicker-textured, and the outer may also be somewhat expanded rather than awl-shaped.Around Tulbagh, where the two spe cies co-occur, O. thyrsoides favours moister, loamy soils while O. dubium prefers drier, stonier situations.Carefiil examination o f mixed populations has failed to reveal a single hybrid plant.In the Warm Bokkeveld around Ceres, O. thyrsoides is replaced by O. ceresianum.Although vegetatively similar and occupying similar habitats, O. ceresianum is recognized by the very large, dark centre to the flower, covering the lower half of the tepals, and by having all six filaments winged, not just the inner three.
The species is rather variable in the shape o f the inflor escence, which may range from narrowly cylindrical to subcorymbose, and in the size o f the flowers.Many of these forms were accorded species status by Leighton (1944) but we prefer to treat them as part of the nor mal variation of the species.Among the more distinc tive local forms are those from Langebaan and Saldanha {Boucher 6476, 6477\ Craven 175), which are anoma lous in their dark, brownish tunics.A particularly attrac tive form, with dense, subcorymbose racemes of flowers with distinctive, brownish centres occurs around Darling and Saldanha.In the Olifants River Valley some individ uals have unusually long, narrowly cylindrical racemes (e.g.Boucher 2617), thus resembling O. strictum.The dark ovaries or bases to the tepals, strongly winged inner filaments, and erect styles, however, are more consistent with O. thyrsoides.Further north, populations between Klawer and Kamieskroon are robust, with arcuate, rather woody pedicels in fruit.Finally, Barker 3332 from Aries Kraal near Elgin includes plants with unusually short styles and scarcely winged inner filaments.Although resembling O. dubium in these features, the long, pointed leaves and small bulbs of these plants are more consis tent with O. thyrsoides.It is possible that occasional hybridization may be the origin of some of these anoma lous plants.
History: Ornithogalum thyrsoides was among the ear liest species in the genus to be described from southern Africa and the first in section Aspasia.The winged inner filaments led Baker (1897) to conflate the species with O. dubium, and even Obermeyer (1978) was misled into treating O. gilgianum from Bainskloof (here included in O. dubium) as a synonym despite its dark tunics.Since the revision of Leighton (1944), however, the species has mostly been correctly understood.
Plants 200-350 mm high.Bulb subglobose, (11-)13-20 mm diam., outer tunics whitish, sometimes brown ish grey, somewhat leathery or membranous.Leaves 3-7, suberect, more than half as long as inflorescence, sometimes reaching to base of raceme, partially or mostly dry at flowering, oblong-lanceolate, 40-90(120) Distribution and ecology, a local endemic o f the Warm Bokkeveld around the town of Ceres (Figure 10), where it occurs in loamy soils in seasonally moist vleis and along seasonal streams, usually in dense colonies.
Discussion: although vegetatively similar to Ornitho galum thyrsoides in its small bulb with pale, membra nous tunics and slender, suberect leaves clasping the stem at the base, O. ceresianum is distinguished by the large, dark olive central eye that covers the basal third o f the tepals, the glossy black ovary, and by having all six filaments expanded and winged in the lower two thirds.The inner filaments are more broadly winged than the outer and in some individuals may be petaloid with a dark base.In O. thyrsoides the outer filaments are at most slightly expanded, the dark centre of the flower is much smaller, covering no more than the base of the tepals, and the ovary is ovoid and matt.In addition the racemes are typically longer, with the lower pedicels shorter in fruit.
This striking species deserves to be in cultivation, and there is little doubt that it can be as easily grown as Ornithogalum thyrsoides.
History: first collected in the early years o f the twen tieth century, Ornithogalum ceresianum was described some twenty years later by Frances Leighton (1933a) but was subsequently treated as merely an unusual form o f O. thyrsoides (Obermeyer 1978).Having seen the taxon in the wild, however, we are convinced that it is sufficiently distinct from O. thyrsoides in several respects to warrant recognition as a distinct species.

Series 3 M aculata
Outer bulb tunics often dark; flowers white, yellow or orange, with or without dark centre; seeds 0.5-1.0mm long.
Flowering time: September-November at lower altitudes but December-January at higher altitudes.Figures IG, 12.
Distribution and ecology: widespread through the southwestern and southern Cape, from the Bokkeveld escarpment southwards to the Cape Peninsula and thence eastwards along the coast and through the Little Karoo as far as Kentani and Stutterheim in the Eastern Cape (Figure 13).The species typically grows on stony flats or rocky slopes in pockets of shallow soil or humus in rock crevices, sometimes along rocky stream banks, in a wide variety of soils, from fine-grained clay to limestone or sandstone and in a range of vegetation types, including renosterveld and fynbos.Discussion: as circumscribed here, Omithogalum dubium is defined by the firm-textured, dark brown to black bulb tunics, spreading, rather blunt leaves, seldom up to half as long as the infloresence and typically densely ciliate on the margin, and corymbose to subcorymbose raceme of white, yellow or orange flowers, usually with a dark centre.The filaments are at least partially concolorous with the tepals, often with a dark zone in the middle or upper part, and the dark-coloured ovary typically narrows into the style, which may be shorter than or subequal to the ovary.The seeds are minute, 0.5-1.0mm long, and echinulate.Similar small seeds are characteristic of all members of series Maculata.
Yellow-or orange-flowered plants with dark outer bulb tunics and ciliate leaves are readily assigned to Ornithogalum dubium but white-flowered individuals may pose some problems.The dark tunics, relatively short, spreading leaves and corymbose inflorescence serve to distinguish such plants ft-om O. thyrsoides and O. strictum, which also have larger seeds, 1.5-2 mm long.O. pniinosum, fi-om fiirther north in Namaqualand, has similar dark tunics and small seeds but the leaves are distinctly glacous and usually glabrous or minutely puberulous on the margins.O. corticatum from the Roggeveld Escarpment is distinguished by its thick, almost corky bulb tunics, leaves that are dis tinctly pubescent on the upper surface near the ends, plain white tepals and filaments, and yellow ovary.On the Cape Peninsula, O. conicum is recognized by its large bulb and linear-filiform filaments and larger seeds.In the Eastern Cape, white-flowered O. dubium is distinguished from O. synanthifolium by its deciduous habit, shorter leaves with ciliate margins, subcorymbose inflorescence and much smaller seeds (0.5-1.0 mm vs 2-3 mm).
Omithogalum dubium is easily the most variable spe cies in the section, displaying almost the entire range of flower colours and filament forms found among the spe cies of sect.Aspasia.Populations are often constant in flower colour, comprising plants that are either white-flow ered, or yellow-to orange-flowered, sometimes with less than one kilometre separating populations of different colours.However, varicoloured populations are known from throughout the range of the species.We have seen a good example of such a population at Hottentotskloof, east of Ceres, comprising individuals displaying the entire range of different flower colours, fi-om orange and yellow through cream-coloured to white with brownish centres.
Plants that match the type of Omithogalum dubium and from which the original collections are most likely to have been made, occur around the town o f Tulbagh at the head of the Breede River, on the original track into the interior.These populations have bright yellow or orange flowers and orange filaments, the inner expanded at the base and apically pointed but not winged, and short styles.Populations from the Swartland west of Tulbagh in contrast, are white-flowered with brownish orange centres.The filaments in these plants are darkly marked in the middle, and the inner are slightly winged.These populations often co-occur with O. thyrsoides, which is common on the coastal side of the mountains, but hybrids between them have not been seen.

Further down the Breede River Valley, between
Worcester and Robertson, populations may be either whiteor orange-flowered, and the inflorescences, although usu ally corymbose, may be longer and more conical.In some of these plants all six filaments are winged, the outer fila ments with acute, divergent wings and the inner with lon ger, convergent wings.Both filament whorls are often stained dark in the middle or in the upper parts.A simi lar degree of variation is evident in the Little Karroo and southern Cape, where populations may be white-or yel low-, or sometimes orange-flowered, and the filaments vary from almost linear to winged.Although white-flowered, longer-styled plants typi cally occur on sandstone soils throughout the moun tains o f the southwestern Cape, and short-styled plants on inter-montane shale soils east of the Bokkeveld-Hottentots Holland Mountain axis, this ecologicalgeographical differentiation is not absolute.Yellowand orange-flowered plants occur, for instance, along the Tsitsikamma Mountains and in the Longkloof on sandstone substrates.Plants from the Witteberg near Laingsburg, including the type of Ornithogalum fimbrimarginatum, are long-styled but otherwise match typical O. dubium perfectly in their dark centre, blackish green ovary, and filaments with the outer portion flushed olivegreen.Apart from the length of the style there is nothing to separate these plants from white-flowered O. dubium.
It is now apparent that there are all transitions between short-styled and longer-styled plants, and that style length alone is an insufficient basis for distinguishing between species in the O. dubium complex.In the light of the extensive variation in flower colour and filament morphology that is now known to occur within even O. dubium sensu stricto, we therefore prefer to recognize a single, variable species in the complex pending more thorough investigation into the population genetics in the group.
The status o f Ornithogalum subcoriaceum in particu lar deserves further study.This species was recognized for small plants, rarely up to 200 mm tall, from the inte rior mountains of the West Coast, typically bearing rather small flowers with tepals 8-14 mm long and bracts that are ciliate towards the apex.In addition, the seeds of collections that we have been able to examine are papil late rather than echinulate as is typical in O. dubium/0.fimbrimarginatum.These distinctions, however, are not absolute and larger plants from the Cedarberg that are referable to O. dubium/0.fimbrimarginatum on this basis may also have ciliate bracts and papillate seeds.We thus provisionally include the smaller plants within a broadly circumscribed O. dubium.
Ornithogalum dubium, unlike O. thyrsoides, has been shown to be non-toxic (Obermeyer 1978).All o f these were treated as varieties of a very broadly defined O. thyrsoides by Baker (1897).The first modem treatment o f the species is that o f Leighton (1944) but the high degree o f variation in flower colour and fila ment morphology evident in this widespread species led her to distinguish almost a dozen different taxa, based on small differences in the shape and size of the filaments and length of the style.Many of these species were reduced to synonomy under O. dubium by Obermeyer (1978), who defined the species more broadly to include all plants with dark bulb tunics, ciliate leaves, and yel low, orange or white flowers with a dark centre and short style.Plants with dark tunics and white flowers with lon ger styles were excluded from this circumscription and treated as separate species, either O. subcoriaceum or O. fimbrimarginatum.
Ornithogalum subcoriaceum was described by Louisa Bolus in 1934 from small plants collected on the Bokkeveld Escarpment near Nieuwoudtville that had ciliate bracts near the apex.Although maintaining the species in her treatment of the genus, Obermeyer (1978) recognized its similarity to O. dubium, and sug gested that the dwarfing might be the result of the colder climate at high altitudes.Plants that match the type of O. subcoriaceum occur throughout the mountains of the southwestern Cape inland of the West Coast, from near Nieuwoudtville to Ceres.Ornithogalum fimbrimatginatum, described by Leighton in 1944 from plants col lected at Whitehill near Matjiesfontein, has never been well understood, and Obermeyer (1978) even considered the possibility that it was a hybrid between O. dubium and O. conicum, inheriting the dark tunics and expanded inner filaments from the former, and the white flowers and long style from the latter.
Ornithogalum dubium was separated from O. fimbri marginatum and O. subcoriaceum by Leighton (1944) and Obermeyer (1978) on the length of the style, which was taken to be less than 2 mm long in O. dubium and more than 2 mm long in 0.fimbrima?ginatumand O. subcoria ceum.Long-styled plants with dark tunics and white flow ers are scattered throughout the mountains of the south western and southern Cape, typically growing in moist, shallow soils in sandstone outcrops.Populations with the outer filaments variously broadened were referred by Leighton (1944) Obermeyer (1978), despite the fact that the length of their styles exceeds the upper limit accepted by her for that species.
Distribution and ecology'-, widespread through Namaqualand, especially along the western edge of the escarp ment between 500-1200 m, from the Richtersveld in the north to Klawer in the south, and extending eastwards to Calvinia on the Bokkeveld Plateau and Aggenys in Bushmanland, with two records from as far east as Kuruman and Augrabies in Gordonia (Figure 15).In the south of its range, in the Knersvlakte, the species occurs along the western foothills of the escarpment below the Bokkeveld plateau, and only extends onto the plateau itself, around Calvinia, from the more arid val leys of southern Bushmanland.Plants typically grow on dry, open stony flats and slopes, and on rocky hillsides, mostly in clay and granite.
Discussion: Ornithogalum pruinosum is recognized by its dark, blackish outer bulb tunics, characteristic glaucous, leathery, often undulate leaves that are seldom up to half as long as the inflorescence, and a subcorym bose inflorescence of pure white flowers (rarely with a small dark centre).The yellow ovary and style in many plants is also diagnostic.The small seeds ± 1 mm long, and dark, leathery tunics place the species in the group that includes O. dubium, O. puberulum, O. maculatum and O. rupestre.Although centred south o f the range of Ornithogalum pruinosum, in the southwestern Cape south of Klawer, outlying populations of O. thyrsoides have been recorded from the higher-lying parts of central Namaqualand, in the Kamiesberg and near Steinkopf.The two cannot be confijsed, however, as O. thyrsoides is readily distin guished from O. pruinosum by the slender, bright green leaves, small bulb with pale tunics, and inner filaments that are conspicuously winged in the lower half and par tially obscuring the ovary.In the south, the distribution of O. pruinosum also approaches that of O. strictum and O. dubium but the species do not overlap and conftision between them is unlikely.O. strictum, which is common in seasonally damp sites along the Bokkeveld Mountains, has similar flowers but slender, bright green leaves and cylindrical racemes with short pedicels, and O. dubium, which has been recorded from similar habitats on the Gifl^erg, has bright green leaves that are densely ciliate on the margins.
Ornithogalum pruinosum is very variable in stature, ranging in size from small, few-flowered plants scarcely 50 mm high to robust individuals almost 700 mm tall.Plants from Aggenys in the arid eastern Bushmanland are especially dwarfed.This variation is to be expected in a species from arid environments with variable pre cipitation.Although usually lanceolate, the inner fila ments are occasionally expanded at the base, the expan sions with small, pointed apical wings.Plants with these winged filaments are scattered through the range of the species, and are known from the Richtersveld, central Namaqualand and Loeriesfontein.The range in form of the inner filaments in the species was evident to Leighton (1944), who noted that they varied in the same popula tion or individual, even in the same flower.
History: Ornithogalum pruinosum appears to have been first collected in the late 19th century, first near Kamieskroon by R. Schlechter in the spring of 1897 and a year later at Okiep by W. Morris.Although Schlechter intended describing the species under the name O. glau cophyllum, it was only formally recognized in 1944, when it was independently described by both Leighton and Von Poellnitz, the latter using Schlechter's material and his proposed epithet.Since then it has been consis tently recognized, and its characteristic glaucous, often undulate leaves have spared it the further taxonomic fragmentation on the basis of the shape of the inner fila ments that has been the fate of other species from the southwestern Cape.
Distribution and ecology': localized to the higherlying parts o f central Namaqualand between Springbok and Garies, but widely scattered through the drier parts of the southwestern Cape, from the Bokkeveld Plateau southwards through the Cedarberg and Cold Bokkeveld to Langebaan and Malmesbury on the west coast, thence eastwards along the fringe of the Cape Floral Region to Prince Alfred (Figure 17).Plants are typically restricted to rock outcrops, typically in shallow humus on granite or sandstone pavements, but also outcroppings o f shale.
Discussion: Ornithogalum maculatum is one of the most striking species in section Aspasia, readily recog nized by its glaucous leaves lacking any marginal vestiture, orange or yellow flowers, usually with some dark marking at the tips of the outer tepals, and subulate or lanceolate filaments.Forms with pure yellow or orange flowers may be confused with O. rupestre but that spe cies has narrow, subterete leaves.The minute seeds and orange flowers might lead to confusion with forms of O. dubium, but O. maculatum lacks the dark tunics and ciliate leaves that are characteristic of that species.
Ornithogalum maculatum is very variable in stature, flower size and in the development of markings on the flowers.Large forms, with pure orange flowers lack ing markings and with broad filaments, originally seg regated as O. splendens, occur around Calvinia and in Namaqualand between Kamieskroon and Hondeklipbaai.Plants of similar size but with well-marked outer tepals bearing triangular or diamond-shaped markings were distinguished as O. speciosum and have been recorded from Namaqualand, between Springbok and Garies, from the Olifants River Valley around Clanwilliam, and from the western Little Karoo, around Karoo Poort, Touws River and Montagu.
The presence and degree of marking may vary within a population, which can thus include both marked and unmarked individuals (e.g.Van der Menve 246).Although typically blackish, the markings on the outer tepals range in colour from green to black, sometimes within a single population (e.g.Horrocks 12; Hall s.n.NBG68697).Occasional plants or populations from Karoopoort (Viviers 1568), Laingsburg (Bayliss 736) and near Steytlerville may lack yellow or orange pigment, producing whitish flowers with darker markings.Such plants formed the basis of O. rossouwii.
Anecdotal evidence suggests that the species is not toxic to animals since goats have been recorded as con suming it with impunity (Obermeyer 1978).
History: first described in 1791 by Nicolaas von Jacquin from plants grown in Vienna, Ornithogalum maculatum was redescribed a few years later by Thunberg from wild material gathered near Saldanha.Struck by the distinctive marks on the outer tepals, he chose the same epithet for his species.The species is now known to vary not only in flower colour but also in the degree of maculation.Some o f the more distinc tive of these forms were segregated as distinct spe cies or varieties in the past.Well-marked forms from Namaqualand were segregated as O. speciosum by Baker (1891), and an attractive form with large, unmarked yel low or orange flowers from near Nieuwoudtville was named O.splendens by Bolus (1931).Both o f these forms were later reduced to varietal status by Leighton (1944) but we are in accordance with Obermeyer (1978) in preferring to regard them as local forms within a much greater spectrum of variation and thus not deserv ing of taxonomic rank.Most recently, Miiller-Doblies & Muller-Doblies (1996) described O. rossouwii from five greenhouse-grown plants collected between Laingsburg and Janseville, and distinguished from O. macula tum by their small size and whitish or pinkish flowers.Although the type material of this species has not been deposited in any herbaria, the accompanying illustra tions make its identity clear.The differences between this material and typical O. maculatum   Plants (30-)50-150(-200) mm high.Bulb subglobose, (7-)10-15 mm diam., outer tunics whitish or brownish.Leaves (2)3-10, subterete or canaliculate, 20-110 x 1-3 mm, glabrous.Raceme corymbose to subcorymbose, 5-30 mm long at flowering, elongating to 10-50 mm in fruit, (l)2-10(-15)-flowered; lowermost pedicels 2-18 mm long, elongating to 6-25 mm in fruit; bracts ovate, whitish with upper part brownish, acute or acuminate.usually shorter than pedicels but longer in small plants, lowermost (4-)6-15 mm long.Flowers usually yellow or orange, rarely white or pinkish; outer tepals ovatelanceolate, inner obovate, 6-12 x 3.5-5.5 mm.Stamens ± half as long as tepals; filaments 3-5 mm long, whitish or yellowish, outer linear to subulate, inner lanceolate.Ovary' ovoid, 3-4 mm long, yellowish; style short, erect, 1-2 mm long.Capsule fusiform to oblong-ovoid, 10 mm long, 3-lobed, apiculate.Seeds angular-pyriform or comma-shaped, 0.75 mm long, rugulose.Chromosomes'.2n = 12 (Pienaar 1963), 20 (Johnson & Brandham 1996).
Flowering time: late August to early October.Figures 1J,  18.
Distribution and ecology': widely distributed through Namaqualand and the more arid southwestern Cape, from the Richtersveld in the north through the higherlying parts of Namaqualand to the Bokkeveld Plateau, southwards through the northern Cedarberg to Darling in the southwest, and through the Cold Bokkeveld to Barrydale and Laingsburg in the southeast, with isolated records from the Roggeveld Escarpment (Figure 19).Plants grow in shallow humus and soil on rock sheets, often on granite or sandstone, where they may occur in large numbers fringing depressions on the rocks that accumulate water in the rainy season.
Discussion: Ornithogalum rupestre is one of the most recognizable species in the section, identified by the few to many, subterete, glabrous leaves.Other spe cies in section Aspasia have plane, oblong to lanceo late leaves.It is typically less than 100 mm high, with pure yellow or orange flowers with linear to lanceolate filaments and a very short style.O. rupestre shares gla brous leaves, mostly yellow or orange flowers with a rudimentary style, and minute, rugulose seeds with O. maculatum and it is likely that the two are immediately related.The flowers of O. rupestre are typically bright yellow or orange but populations from Langebaan and the Cold Bokkeveld have pale yellow flowers, and plants with cream-coloured flowers have been collected around Pakhuis Pass.The flowers of populations from the Vredenburg Peninsula are uniformly white or pinkish.
History: Ornithogalum rupestre was described by Linneaus f (1782) from plants collected on Witteklip near Vredenburg by Carl Peter Thunberg.Although not mentioned in the original description, the plants from Witteklip are uniformly white-flowered, sometimes with a pinkish flush.Later collections of the more common and widespread yellow-flowered forms were described as O. multifolium, based on short, floriferous plants from Namaqualand (Baker 1873), and O. aurantiacum, from few-flowered plants collected near Mamre (Baker 1897).Yet another name, O. ranunculoides, was applied to taller forms from Namaqualand by Louisa Bolus (1933b).All of these forms were subsequently included within a single yellow-flowered taxon, O. multifo lium, by both Obermeyer (1978) and Muller-Doblies & Miiller-Doblies (1996).White-flowered O. rupestre from Vredenburg has until now been retained as distinct from yellow-flowered O. multifolium.The application o f the name O. rupestre was confused by Leighton (1944), who used it for yellow-flowered plants (Obermeyer 1978), but there is no doubt that the type is white-flowered.
White-flowered Ornithogalum rupestre was separated from yellow-flowered O. multifolium on nothing more than flower colour, a minute difference in style length, and the smaller stature of the former.It is now clear that flower colour varies within the species in the group and is an insufficient basis for separating species.Even within yellow-flowered plants, flower colour may range from pale lemon-yellow through bright canary yellow to orange, whereas paler forms may be white, creamcoloured or pinkish.The purported differences in stat ure and style length are likewise trivial and we have no hesitation in including the various colour forms within a single species.It is unfortunate that O. rupestre is the earlier name for this taxon, which is better known under the name O. multifolium.

V < 3 m
Discussion: still poorly collected and understood, Ornithogalum leeupoortense is distinguished by the fan like arrangement of its leaves, which are falcate, pointed and somewhat channelled.The leaf margins are obscurely papillate and in the populations around Springbok they are also distinctly thickened.Plants typically produce a second inflorescence in a growing season and the older inflorescence is characteristically displaced, appearing to arise in the axil of one of the lower leaves.The relatively short inflorescence is characterized by large, often foliose, green bracts, the lowermost 15-25 mm long, with the tips drawn out and attenuate.
Collections from the Richtersveld, although lack ing the thickened leaf margins of plants from around Springbok, match the southern populations in all other essentials, including the falcate, glossy green leaves with attenuate tips, and lateral displacement of the pri mary inflorescence through development of a later one, and we have no hesitation in regarding them as conspecific.Distribution and ecology, largely restricted to a small area of high ground in southern Namibia, between 600-950 m, in the Klinghardt Mountains and nearby Huib Hoch Plateau (Figure 22), with a southward extension into the Richtersveld in Northern Cape, South Africa around Lekkersing.This is an extremely arid region that is summer-dry, although coastal fogs supply some pre cipitation in the autumn.
The leaves of Ornithogalum pubendum are remark ably soft-textured for a species from such an arid area and plants are restricted to rock outcrops and crevices, on south-facing exposures or at the foot of boulders, where they are sheltered from the afternoon sun in win ter and benefit from the cooler, moist conditions during the growing season.The species has been recorded as locally common where it occurs but leafing and flower ing is dependent on adequate rain.Discussion: this highly distinctive species is immedi ately recognizable by the two (rarely three) soft-textured, ovate to elliptical (rarely lanceolate) leaves that are either glabrous or pubescent but always with soft cilia 0.7-2.0mm long on the margins.The leaves of Ornithogalum pubendum are typically rather tubular and inflated at the base where they clasp the stem, and the upper leaf is mostly only half the size of the lower one or two.
The species shows considerable variation in the pubescence of the leaves, sometimes within populations but more usually between them.The type collection comprises four plants, all of which have leaves that are entirely pubescent on both surfaces, apart from the upper leaf on one o f the plants, which is pubescent only apically.(1996).Although a similar range in inflorescence form is shown by other species in the group, including O. p minosum, the status of these populations requires fijrther investigation.
The characteristic leaves of Ornithogalum pubenilum, clasping at the base and with long-ciliate margins, have led to confiision with O. hispidum in the herbarium.The latter species, a member of section Hispidaspasia, is readily distinguished by its small bracts, and narrow raceme of smaller flowers with filiform filaments.History'' .Ornithogalum puberulum was described by Obermeyer (1978) from plants collected fifteen years earlier in southern Namibia.Ornithogalum merxmuelleri, described the following year by Roessler (1979) from fruiting material collected in the same area of southern Namibia, does not appear to differ in any sig nificant way and was synonymized under O. puberulum by MUller-Doblies & Muller-Doblies (1996).
At the same time, Muller-Doblies & Muller-Doblies (1996) described O. puberulum subsp.chris-bayeri from plants that they had collected near Rosh Pinah in south ern Namibia, distinguishing it by the shorter cilia on the leaf margins, hairy bracts, and yellow-spotted filaments and short style.These differences are, however, not sig nificant in the light of more recent collections.We owe much of our knowledge of the species to the recent col lections made by the Cape Town succulent specialist, P.V. Bruyns.

Excluded species
1. Ornithogalum baurii Baker in Flora capensis 6: 504 (1897).Type: South Africa, Eastern Cape [Transkei], Baziya Mountain, 4000' [1 220 m], November without year, Baur 552 (K, holo.!; SAM, iso.!).These three taxa comprise a group o f poorly under stood species known fi-om isolated localities along the mountains of the Eastern Cape and KwaZulu-Natal.They are all small or dwarf plants with two or three leaves and subcorymbose racemes o f pure white flowers subtended by foliar bracts.The seeds of Ornithogalum diphyllum are club-shaped and echinulate (Obermeyer 1978) and those of O. sephtonii tetragonal and reticulate (Hilliard & Burtt 1983).In the light of this, Obermeyer (1978) and Muller-Doblies & Muller-Doblies (1996) had little hesitation in associating this group of species with section Aspasia from the Western and Eastern Cape, despite the obvious anomaly in their distribution.They were interpreted by Obermeyer (1978) and others as iso lated montane relics of the Cape section Aspasia.
At the same time Obermeyer (1978) included O. bau rii and O. synanthifolium, both o f which had been col lected by Baur at Baziya in Eastern Cape, in a broadly circumscribed O. conicum.There is little doubt, how ever, that O. baurii and O. synanthifolium are distinct species (see Discussion under the latter), an opinion that was first aired by Hilliard & Burtt (1988, 1989).What is not as evident, however, is where the affinities of O. baurii, O. diphyllum and O. sephtonii lie.While there is certainly a superficial similarity in vegetative and floral morphology between these three species and those of section Aspasia, particularly in their foliar bracts and moderate-sized, white flowers, and while the seeds of O. diphyllum at least have some resemblance to those of other species in the section, the capsules of this group are not at all like those that characterize section Aspasia.All three species have distinctly turbinate, 3-angled capsules that are exposed by the reflexing of the tepals.Such capsules are anomalous in section Aspasia, which is characterized by ellipsoid or ovoid, obscurely angled capsules that are concealed by the persistent, papery perianth.On this basis alone the three species of the O. baurii group should be excluded from section Aspasia.A F e l l l n s h M D«i Support for this interpretation is provided by DNA sequence data (Manning et aL in prep.), which places O. sephtonii among the residue of species previously grouped within subgenus Urophyllon, where such turbi nate capsules are common, and not among the species of subgenus Aspasia.We accordingly exclude these three species from section Aspasia.The differences between them are relatively minor and further study is likely to reduce the number o f taxa.are also grateful to the curators of the various herbaria for access to their collections.Clare Archer is thanked for her assistance with specific queries.Additional mate rial was collected under permits from the Northern and the Western Cape Nature Conservation authorities.
IO N A L BO TA N IC G A R D E N S OF SO U TH A F R IC A , K iw rfWOtCM.w m jL H om, c.m.
Yellow-flowered populations are known from south of the Riviersonderend Mountains, between Caledon and Bredasdorp, and formed the basis of Omithogalum citrinum, described by Von Poellnitz from plants collected by Rudolph Schlechter at Genadendal in the southern foothills of the Riviersonderend Mountains.Plants from higher altitudes throughout the mountains o f the south western and southern Cape are invariably white-flow ered, typically with longer styles, and were previously segregated asO.fimbhmarginatum and O. subcoriaceum.
History.Ornithogahim dubium, described by Houttyn in 1780, was the second member of section Aspasia known to science after O. thyrsoides.Other yellow-and orange-flowered plants formed the basis o f Jacquin's (1789b)O.flavescens,O. flavissimum and O. miniatum.
are trivial when compared to the variation within the species.Several collections of depauperate plants with the yellow, spot ted flowers of typical O. maculatum (e.g.Pretorius 92) are indistinguishable from O. rossouwii apart from the colour of the flowers, and these white or cream-coloured plants are most appropriately treated as colour forms of O. maculatum.A similar range in flower colour is com-i t h o ^l u * a a e u l i t u * J a c q .e u 1 a t u a H o r t h t r n B o k k « v t l d : f l a t s a t w e « t b a s e o f Q e e l b s r Ŝ * i i d y o p « n f l a t * : B h n l l o w s ^o d o n r o c k a u r f a c * * .t w o g r o u p s o f p l a n t s ; f l o t * * r « o r a n c # w i t h b l a c k p a t c h o n r s v « r s « o f o u t s r a s c a t n t s .
l t h o f * l u « p u b * r u l u « O b # r « .S p « r r g » b t » t .H i l U SE o f H e i o « b t o w a r d * U g u c h a b R i v e r .S t o n y SM s l o p « « i .k g r o u p * o f * o f t L L e a v e s j r e l l o w l s h g r * * n F l o w a m w h i t e .

lanceolate to oblong, usually glabrous
. Type: O. conicum Jacq.filaments filiform to lanceolate and all similar or inner broader, both whorls or more usually only inner vari-
, included not only plants from the Eastern Cape with narrow filaments, which are segregated here as O. synanthifolium, but also collections from the Olifants River Valley that are treated here as a distinct species O. strictum.Both of these taxa have relatively long leaves that are still green at flowering, leading Louisa Bolus to describe O. aestivum for plants from the West Coast that produced a rosette of short leaves which were dry and withered by the time that the plants flowered in summer.
The distinction between the two is not always clear, however, especially in the Olifants River Valley, where occasional plants with the narrowly cylindrical raceme of Ornithogalum strictum have winged inner filaments like those of O. thyrsoides.A similar situation arises further north in Namaqualand.An alternative treatment would be to regard O. strictum as the northern subspecies of O. thyrsoides but intermediate types are rare and we prefer to maintain the two taxa as separate species, especially in view of the unusual declinate style that appears to char acterize O. strictum.History: described by Louisa Bolus in 1933 from plants collected on Vanrhyn's Pass on the Bokkeveld Escarpment, Ornithogalum strictum was included in O. conicum by subsequent authors

glaucophyllum Schltr. ex Poelln.: 23 (1944). Type: South Africa, [Northern Cape], Arakup [Arkoep, N o f Kamieskroon], 14 September 1897, R. Schlechter 11249 (Breslau, holo.; BOL!, GRA. K!, PRE, iso.).
Obermeyer (1978)ants collected by Rudolph Schlechter on 10 December 1896 near Elim in the Overberg.Obermeyer (1978)associated these plants with a collection of O. dubium made in the vicinity on the same day by Harry Bolus {Bolus 8696), who was collect ing with Schlechter at the time, but Miiller-Doblies & Miiller-Doblies (1996) treated O. perpulchrum as a syno nym o f O. rupestre (as O. multifjlium) on the basis that duplicates of both Schlechter 9681 and Bolus 8696 that they examined had the leaves o f O. multifi)lium.This is, however, most definitely not the case with the Bolus col lections in both the Bolus and Kew herbaria, which have the short, oblong, ciliate leaves and dark tunics of O. dubium.Similarly, we have no hesitation in identifying the Kew isotype of Schlechter's collection as O. dubium, despite the rather small stature of the plants.There is no doubt that Schlechter's Rietfonteinkloof is correctly located near Elim (Jessop 1964), and Ornithogalum ntpestre (= O. multifolium) is not known from south of the Langeberg, whereas O. dubium is common here.All evidence thus confirms that Ornithogalum perpulchrum is correctly placed in synonomy under O. dubium. O.

merxmiielleri Roessler: 389 (1979). Type: Namibia, Farm Namuskluft, 29 September 1977, Merxmiiller cfe Giess 32350 (M, holo.; PRE!, WIND, iso.). O. piibenilum subsp. chris-bayeri U.M iill.-Doblies & D.Mull.- Doblies: 390 (1996), syn. nov. Type: Namibia, mountain 2 km west o f Rosh Pinah, ± 450 m, 26 September 1989, M iiller-Doblies 89120b (WIND, holo., not deposited).
The minute, echinulate seeds o f Ornithogalum leeupoortense indicate an alliance with the members of the O. dubium group but it is unlikely to be confused with any other species in series Maculata on account of the distinctive glossy green, distichous leaves.Most other species o f the group from Namaqualand, particularly O. pruinosum and O. maculatum, have rosulate, glaucous leaves.Pressed specimens of O. pruinosum may be dis tinguished by their erect, proportionally broader leaves with ciliolate, often crisped margins.The leaves are also typically shorter, less than half as long as the inflores cence, and thick-textured.History'-, the species was described by Muller-Doblies & Muller-Doblies (1996) from a collection of non flowering plants made on 11 August 1988 on the Farm Leeupoort northeast of Springbok and subsequently grown on in the glasshouse.None o f the cited material has been deposited in any herbarium but three collec tions of plants, all from Naip se Berg on the edge of Bushmanland northeast of Springbok, match the protologue in the diagnostic essentials, viz. the distichous leaves with thickened, obscurely papillate margins, and are accordingly referred to this species.Since no type material has been lodged in the cited herbaria, we select one of these specimens to act as a neotype.The earliest collection of the species that we have traced is that made by Margaret Thompson and Annelise le Roux in August 1977 in the Richtersveld and until now referred either to O. subcoriaecum (now a synonym of O. dubium) or O. pruinosum.Raceme corymbose or racemose, 15-100(-130) mm long at flowering, 4-10(-25)-flowered; lower most pedicels 10-20 mm long in flower, elongating to 30 mm; bracts whitish, ovate, acute or acuminate, glabrous or lowermost ciliate, shorter or longer than pedicels, lowermost 8-16 mm long.Flowers white; outer tepals O.

FIGURE 21.-Known distribution o f Ornithogahm leeupoortem e, O; O. pubendum , • . have
The leaf and bract margins in all four plants are densely ciliate.The type of Ornithogalum merxmuelleri has similarly pubescent leaves but the bracts are entirely glabrous.All other collections that we have examined have leaves that are glabrous with sparsely pubescent margins.Although both the type and Bruyns 2668 have ciliate margins to the bracts, other populations from near Rosh Pinah {Bruyns 8359, 8866) have entirely gla brous bracts.Described from near Rosh Pinah in southern Nami bia, Ornithogalum puberulum subsp.chris-bayeri Mull.-Doblies & Miill.-Doblies was distinguished from the typical subspecies by the shorter cilia on the leaf mar gins, hairy bracts, yellow-spotted filaments and shorter style (1.6 mm vs 2.5-3.0 mm).More recent collections of O. puberulum, with somewhat variably ciliate leaf margins, glabrous or ciliate bracts and short styles, that subsequently been made near Rosh Pinah {Bruyns 2778, 8359, 8866) indicate that the species is more vari able than was previously supposed and that the recogni tion of this subspecies is unwarranted.Typical Ornithogalum pubentlum from southern Namibia is characterized by a rather corymbose inflores cence with up to 15 flowers.Populations of vegetatively similar plants from near Lekkersing in the Richtersveld {Oliver, Tdlken & Venter 776; Thompson & Le R oilx 85), however, have elongate, cylindrical inflorescences with up to 25 flowers, giving them a very different appear ance.These plants were associated with O. pubenilum without comment by Miiller-Doblies & Muller-Doblies