Systematics of the genus Daubenya ( Hyacinthaceae : Massonieae )

Daubenya Lindl. was until recently thought to comprise the single species D. aurea Lindl. but is now considered to include the monotypic genera Androsiphon Schltr. and Amphisiphon W.F.Barker. as well as the species previously referred to the genus Neobakeria Schltr. Eight species are now recognized in the genus, including the new combinations Daubenya comata (Burch, ex Baker) J.C.Manning & A.M.van der Merwe and D. zeyheri (Kunth) J.C.Manning & A.M.van der Merwe. Each species is fully described and illustrated in black-and-white and in colour. A key to the species, and distribution maps


INTRODUCTION
The Hyacinthaceae (excluding the North American chlorogaloid genera) are now well established as a monophyletic lineage within the order Asparagales (Fay & Chase 1996;Pfosser & Speta 1999;Fay et al. 2000).The family is distributed through Africa, across most of Europe and central Asia to India, and in Andean South America, with centres of diversity in southern Africa and the Mediterranean (Speta 1998).The plants prefer open.sunny habitats in seasonal climates and are correspond ingly rare in heavily wooded, tropical regions.Although it has long proved difficult to devise a natural infrafamilial classification of the Hy acinthaceae.recent molecular analysis of the chloroplast DNA region m?L-F provides strong support for the division of the family into four subfamilies, Oziroëoideae, Urgineoideae.Omithogaloideae and Hyacinthoideae (Pfosser & Speta 1999).Within the subfamily Hyacinthoideae two clades are retrieved.The first combines the Indian and African genera south of the Sahara, and largely coincides with the delimitation of the tribe Massonieae Baker (with the significant inclu sion of the sub-Saharan species previously placed in the genus Scilla L.).The second clade includes the Medi terranean and Asian genera, corresponding to the tribe Hyacintheae Dumort (Pfosser & Speta 1999).Despite the strong molecular support for the recognition of these two tribes, few corroborative morphological characters are available.Further division of tribe Massonieae into the subtribes Ledebouriinae and Massoniinae (Muller-Doblies & Muller-Doblies 1997) is not supported by the molecular data.The poor congruence between morpho logical and other characters within Hyacinthaceae has also made generic circumscriptions very difficult.One of the consequences of this has been the recognition of a large number of genera that are poorly defined morpho logically (Speta 1998).
Within the tribe Massonieae the relationships between the species traditionally placed in the genera Amphisiphon W.F. Barker (1 sp.).Androsiphon Schltr.(1 sp.), Daubenya Lindl.(1 sp.).Neobakeria Schltr.(1-3 spp.) and Massonia Houtt.(± 6 spp.) have been especially controversial (Jessop 1976;Muller-Doblies & Muller-Doblies 1997;Goldblatt & Manning 2000).These genera share some morphological characters, in particular two prostrate or spreading leaves and a condensed inflorescence of more or less tubular flowers but differ greatly from one another in floral details.Traditional morphological methods have not been useful in resolving the relationships between the species but molecular analysis has proven extremely in formative.Analysis of chloroplast DNA ( Van der Merwe et al. in prep.)includes Amphisiphon.Androsiphon.Dau benya.Neobakeria and one species of Massonia in a strongly supported monophyletic clade.This clade is resis tant to further segregation at the generic level and the cir-cumscription of the genus Daubenya is therefore broad ened to circumscribe the whole clade.The genus Daubenya in this broader sense is characterized by a tunicated bulb with the older tunics leathery and dark brown and extending as a short papery neck, two ± glossy leaves with impressed longitudinal striations on the upper sur face, and a racemose, corymbose or subspicate inflores cence of white, red or yellow flowers with a slender, cylin drical perianth tube.Species of Massonia s. sir.are char acterized by their soft, fleshy leaves, corymbose inflores cences with large, leaflike bracts, white to pink flowers with the tepals typically sharply recurved from the base and then incurved in a characteristic sigmoid fold, and tumbleweed-like infructescences.
Daubenya is centred along the western edge of the southern African central plateau and all of the species are winter-growing.Most are highly localized endemics restricted to pockets of doleritic clays.The vegetative similarity between the species contrasts sharply with the unusual diversity of floral form evident in the genus.This floral radiation represents adaptations to a variety of pollinating agents, including insects such as bees, butter flies, moths and monkey beetles and sunbirds.Most of the species exhibit adaptations to anemochory, especial ly in the development of large, winged capsules.These winged, dehiscent capsules contrast markedly with the indehiscent, ovoid capsules present in two of the species.Eight species are recognized within this expanded con cept of Daubenya, some of them well established in hor ticulture but others very poorly known.This revision provides full descriptions of all species as well as notes on their history, ecology and distribution.All of the species were studied in the field and each is fully illus trated in black-and-white and in colour.

MATERIALS AND METHODS
Herbarium specimens in BOL, K, NBG, PRE and SAM were studied to gather data on morphology, flow ering time and distribution.Extensive fieldwork was also undertaken and all species were visited in their natural habitat.All observations on pollinators were made in the field.Conservation status for each species was assessed according to the new IUCN categories and criteria (Victor 2000).

MORPHOLOGICAL CHARACTERS OF TAXONOMIC SIGNIFICANCE
Bulb: the bulbs in Daubenya are turbinate to globose or ovoid in shape and between 20-35 mm diam.The outer tunics become leathery and dark brown on drying, and split at the top into narrow, flat, papery segments that form a characteristic neck.This papery neck is particu larly conspicuous in species or individuals with deeply buried bulbs.A similar neck is found in some species of Lachenalia but not in any species of Massonia s. sir.
Leaves: all species have paired leaves that arc pros trate or spreading.The leaf bases are subterranean and long or short depending on the depth of the bulb.The blades are glabrous and rather glossy above with impressed longitudinal striations along the main veins.
They are lanceolate to ovate in shape and measure 40-150 x 25-50 mm.
Leaf anatomy: leaves in Daubenya are amphistomatic with a thick cuticle.The marginal epidermal cells are columnar with slightly thickened walls and a unicellular hypodermis is present at the leaf margin.The mesophyll typically comprises three rows of palisade parenchyma adaxially and three rows of spongy parenchyma abaxially.The vascular bundles are surrounded by parenchyma tous bundle sheaths.Slight indentations correlating with the impressed longitudinal striations on the upper leaf surface occur above the main veins.In these indentations the epidermal cells are smaller than elsewhere with thicker outer periclinal walls and they overlie two rows of palisade parenchyma instead of three.
Inflorescence: is borne at ground level between the leaves on a subterranean peduncle.It is essentially race mose but varies in shape from conical to capitate.Shortening of the rachis in several species results in a corymbose, rather capitate inflorescence, whereas sup pression of the pedicels results in a subspicate inflores cence, accompanied in some species by shortening of the axis.Well-developed pedicels and a corymbose inflores cence are found in Daubenya aurea, D. alba, D. capen sis and D. zeyheri but the remaining species are charac terized by a subspicate inflorescence.This is usually con ical.protruding for up to 90 mm above the leaves in D. marginata and D. namaquensis, but is ± capitate in D. comata and D. stylosa.The inflorescence bracts are usu ally inconspicuous and pale in colour except in D. aurea, which is characterized by large, green bracLs.The bracts typically increase in size acropetally but the degree of this increase varies.In D. comata, D. stylosa and D. zey heri the upper bracts are only slightly larger than the lower ones and form an inconspicuous greenish coma at the top of the inflorescence.In D. namaquensis and D. marginata, however, the increase in bract size is marked and the inflorescences are topped by a more conspicuous coma of greenish or pinkish to orange bracts.The inflo rescence remains at ground level until the fruits are mature, at which stage the peduncle elongates rapidly from the base, pushing the infructescence well above the ground as a prelude to dispersal of the seeds.
Flowers: there is great variation in the morphology of the flowers in Daubenya but all species are characterized by a slender, cylindrical perianth tube.The tepals are usually free and spreading from the level of the stamen insertion but in D. stylosa they are fused well beyond this level into a narrow tube around the staminal column.Tepal colour ranges from white or pinkish in D. alba, D. comata, D. marginata and D. zeyheri, to bright yellow in I), capensis, D. namaquensis and I), stylosa, or brilliant red in most populations of D. aurea.The flowers in most species are actinomorphic but some degree of zygomorphy is evident in the lowermost flowers in D. comata and D. namaquensis, and reaches an extreme degree in D. aurea.In D. comata and D. namaquensis the outer flow ers are at most weakly bilabiate through a slight asym metry in the penanth tube, with the adaxial side slightly longer than the abaxial side.An analagous situation is also sometimes evident in the filament column in the lowermost flowers of D. alba.In D. aurea, however, the zygomorphy is extremely marked and results from the great enlargement of the lowermost tepals of the lower flowers, accompanied by an elongation of the perianth tube on this side rather than on the adaxial side.The sta mens in most species of Daubenya are ± fused at the base into a staminal column or tube.This is most marked in D. alba, D. aurea, D. capensis and D. stylosa.In the bilabi ate, lower flowers of D. aurea, the abaxial tepals are actually fused to the staminal column and as a conse quence of this the abaxial filaments are inserted much higher up the perianth than the adaxial and appear to be free from one another.In D. capensis a thick, convex disc occludes the top of the staminal tube with the style pro truding through a small pore in the disc.The staminal column is very short in D. marginata and D. zeyheri, and is lacking or only slightly evident in D. comata and D. namaquensis.
Fruit and seeds: the fruit in Daubenya is a papery capsule, typically dehiscing loculicidally in the upper portion.In D. comata the capsule is obovoid and round ed but in most of the other species it is ± inflated and three-angled.These angles are particularly well devel oped in the upper part of the capsules in D. alba and D. capensis, and the capsules in these two species are decid edly cuneate in shape.The species D. marginata, D. namaquensis and D. zeyheri are distinctive in their deci duous capsules that fall free of the pedicels at maturity and are dispersed individually.In other species of Daubenya the capsules remain attached to the pedicel and rachis and the entire infructescence functions as the dispersal unit.Unique capsules with a persistent, horn like style characterize D. stylosa.The capsules in this species are essentially indehiscent although the locules eventually separate along the septa.The seeds in all species are globose and black in colour with a smooth testa.They range in size from 2-3 mm diam.

DISTRIBUTION AND ECOLOGY
Daubenya is a small genus of eight species that is endemic to South Africa.It is largely restricted to the fringes of the winter rainfall region along the Bokkeveld and Roggeveld Escarpments, which mark the western margin of the South African central plateau.Five of the eight species occur here, at altitudes of 1 000-1 500 m.Most of the species of Daubenya are highly local endemics and only D. comata is more widespread, occur ring over much of the South African central plateau.All of the species are winter-growing, despite the fact that not all are restricted to the winter rainfall region.Plants typically leaf and flower in the autumn or winter, between April and July, with only one species, D. aurea, flowering in the spring, in early September.Five of the species, D. alba, D. aurea, D. capensis, D. sty losa and D. zeyheri are restricted to the winter rainfall region and respond largely to frontal rain that falls between April and August.Apart from D. zeyheri, which is a coastal species confined to calcareous sands near Saldanha, the remaining four winter rainfall species occur along the edge of the Bokkeveld and Roggeveld Escarpments from Nieuwoudtville and Calvinia in the north southwards to Sutherland.The remaining three species occur partially or wholly within the summer rainfall region.Daubenya marginata is distributed along the Roggeveld Escarp ment but extends inland to Fraserburg, where it enters the summer rainfall region; D. namaquensis occurs at the edge of the summer rainfall region in Bushmanland, east of Springbok; and D. comata is widespread across the central parts of the summer rainfall region.Despite their distribution, these three species respond to autumn thun dershowers and begin to grow slightly before the winter rainfall species.
Most species of Daubenya are highly localized endemics known only from a few populations.They typ ically occur in small colonies, sometimes in large num bers, in heavy clay soils, and are invariably restricted to low-lying washes or drainage lines where the soil be comes seasonally waterlogged.The majority of the species occur on the South African central plateau, where they are largely restricted to clays derived from dolerite.The particular nature of doleritic clays ensures that these soils retain moisture for longer than the surrounding clays derived from shales of the Karoo series.The two remain ing species occur on more sandy substrates, D. namaquen sis in deep red sands and D. zeyheri in calcareous coastal sands.
All of the species of Daubenya are vulnerable to dis turbance or transformation of their habitat, particularly through agriculture.Although predation of the bulbs by porcupines is common among the species growing on the Roggeveld Escarpment, its impact on the populations is unknown.The numbers of these rodents can be expected to have increased as a result of the reduction in their natural predators, particularly leopard.Most species ot Daubenya still appear to set large quantities of seed in the wild to replace the adult plants consumed by herbi vores.An additional pressure on populations comes from sheep and baboon, which eat the leaves and inflores cences, resulting in a drastic reduction in seed set in some instances.A recent initiative aims at the protection of a population of the yellow-flowered form of D. aurea from overgrazing through co-operation of the landowner and the local bulb growers' association.There is, how ever, no formal conservation of this population and its survival thus remains uncertain.In another encouraging development, cultivated bulbs of D. aurea have become available in commercial nurseries.All of the species of Daubenya are worth cultivating for their compact habit and brilliantly coloured, often fragrant flowers.

POLLINATION AND SEED DISPERSAL
The floral diversity evident in the genus Daubenya is exceptional in the family Hyacinthaceae and reflects a corresponding diversity in pollination strategies.These include several strategies that, although well developed in the Western Cape, are rare or absent elsewhere.Most of the species produce fragrant, white or yellow flowers that are visited for nectar or pollen by a variety of diur nal and nocturnal insects, including pollen-and nectarcollecting bees, butterflies and moths in the family Noctuidae.The species with generalist flowers include D. namaquensis, D. comata, D. alba and D. stylosa.In these species the nectar collects either in the perianth tube only or also in the staminal tube above this.Both the perianth and staminal tubes are narrow and the nectar is thus accessible only to long-tongued insects.
Two species, D. marginata and D. zeyheri, are adapt ed to pollination by sunbirds.The adaptations to omithophily in these species include a reduced and widened staminal tube that forms a shallow reservoir in which significant quantities of nectar accumulates, orange or red filaments (with purple bases in D. zeyheri) and a lack of fragrance.Omithophily occurs in several other genera of Hyacinthaceae in the Western Cape, especially Lachenalia and Veltheimia.Daubenya aurea is adapted to pollination by monkey beetles (Scarabidae-Rutellinae), a pollination strategy that appears to be unique among the Hyacinthaceae, although well repre sented among Western Cape Iridaceae.These active, hairy beetles utilize various brightly coloured, open or bowl-shaped flowers as sites for reproduction and are often encountered on various species of Asteraceae and Iridaceae (Goldblatt et al. 1998).Flowers adapted to these beetles are typically ornamented with dark mark ings that act as beetle mimics, decoying the insects through their resemblance to potential mates.Adapta tions for this type of beetle pollination in D. aurea are the asymmetric enlargement of the lower or peripheral (low ers in the inflorescence to imitate the ray florets of Asteraceae like Arctotis and Gazania, the lack of nectar, with associated vestigial staminal tube, and the absence of floral fragrance.Although the pollination biology of the remaining species, D. capensis, has not been studied, the peculiar structure of the flowers and their yeasty odour suggest that the species might be adapted to polli nation by rodents.This strategy is also evident in one or two species of Massonia (Hyacinthaceae) (Johnson et al. 20()1) as well as several species of Androcymbium Willd.(Colchicaceae) among the geophytes in Western and Northern Cape.
In all species of Daubenya the fruiting peduncle elon gates at maturity, raising the infructescence above the ground, and then abscises to release it.Thereafter, the species differ markedly in seed dispersal strategies.In D. comata and D. stylosa no particular adaptations for seed dispersal appear to be developed and the relatively small capsules remain attached to the axis, shedding the seeds immediately around the plants.These autochorous species have short or vestigial pedicels that do not elon gate at maturity and ± ovoid capsules that are indehiscent.although the locules do eventually separate along the septa to release the seeds over time as they disinte grate.The remaining species exhibit a suite of adapta tions to anemogeochory or wind dispersal through tum bling (Van der Pijl 1982;Snijman & Linder 1996).The most prominent of these adaptations are the large, three angled or -winged, loculicidally dehiscent capsule and the frequent detachment of the entire inflorescence as a single unit before seed release.In D. marginata, D. namaquensis and D. zeyheri the capsules abscise readily from the pedicels at maturity and are dispersed individu ally but in the remaining species they remain attached to the peduncle and the infructescence is dispersed as an entire unit.The surface area of the dispersal unit in these species is further increased by the prominent wings on the capsules in D. alba and D. capensis, giving them a kite-like appearance and by the large papery bracts in D. aurea, which act as sails.The conversion of the in fructescence into a tumbleweed is enhanced in D. alba and D. capensis through the elongation of the pedicels, increasing the size of the infructescence and giving it a rounded, balloon-like form.There is a remarkable simi larity in the adaptations to wind dispersal of the fruits developed in these latter species and in the genus Massonia, particularly the obtriangular, winged capsules borne on relatively long pedicels to form a rounded, bal loon-like structure that is readily dispersed by the wind.In Massonia, however, the pedicels are invariably sub tended by large, sail-like bracts resembling those found in D. aurea.The smooth, globose seeds in the anemochorous species are easily shaken out of the capsules but do not appear to disperse significantly thereafter.
Anemogeochory is well known in the family Amaryllidaceae: subtribe Amaryllideae (Snijman & Linder 1996) but has not been studied in the Hyacinthaceae.In both of these families, however, this mode of seed dis persal is more common in South Africa than elsewhere in the continent, and is best developed in the semi-arid, winter rainfall region.

EVOLUTION
Sequence analysis of the tm L-tm F region of the chloroplast genome (Van der Merwe et al. in prep.)offers little resolution within the genus apart from indi cating a close relationship between the species D. alba and D. capensis.This species pair is well defined mor phologically by the cuneate.winged capsules borne on long pedicels.Their relationship to the remaining species as well as the relationships between these species remain unresolved.Although the similarity in floral form and capsules in D. marginata and D. zeyheri suggests that these two species are closely allied, further morphologi cal assessment of relationships within the genus is con founded by the high degree of autapomorphies displayed by each species.Until further genetic analysis is com plete.little more can be said about possible relationships between the species.
The genus Daubenya is essentially restricted to sea sonally moist, usually clay depressions and all of the species respond to autumn rains, whether they occur in the winter or summer rainfall regions.There is thus little ecological diversification within the genus in response to climate or soils, which is probably not surprising given that all species are vegetatively indistinguishable.Apart from D. comata and D. marginata, species of Daubenya are local endemics that are rarely sympatric.Exceptions are the species pairs D. aurea and D. marginata in one or two localities, and D. st\losa and D. capensis.It is per haps significant that the members of these pairs flower at different times, suggesting that flowering time operates as a prezygotic isolating mechanism between them.While there may be little diversification within the genus in response to climate or soil.Daubenya is unparalled in the Hyacinthaceae in its floral radiation.This extreme variation can be linked to differences in pollination strat egy and is accompanied by some variation in capsule morphology and seed dispersal mechanisms.Most species are anemochorous to some degree and adapta tions to wind dispersal of the seeds include both elabora tion of the capsules themselves as well as enlargement of the inflorescence bracts.Exceptions are D. stylosa and possibly D. comata, which have indehiscent capsules with no apparent adaptations for seed dispersal.Although the anemochorous species are more diverse than those lacking obvious seed dispersal mechanisms, the individual species adapted to anemochory are not more widespread than the autochorous species.This sug gests that anemochory in Daubenya has facilitated saltational speciation, whereby the occasional colonization of suitable new habitats through seed dispersal has favoured speciation through founder-effects and subse quent genetic drift.

TAXONOMIC HISTORY
The taxonomic history of the genus Daubenya reflects the significance that has traditionally been attached to floral differences among the Hyacinthaceae, resulting in the establishment of several monotypic genera between 1835 and 1936 for newly discovered species with dis tinctive flowers.The first of these genera, Daubenya, was established in 1835 by Lindley to accommodate the species D. aurea, which is characterized by extremely zygomorphic lower flowers.Floral zygomorphy is uncommon in the family Hyacinthaceae and this degree of zygomorphy does not occur elsewhere in the family.In his treatment of the sympetalous genera of the family Hyacinthaceae, Baker (1871) allied Daubenya with the genus Massonia in his tribe Massonieae on the basis of the reduced peduncle.Bentham (1883), however, includ ed these two genera in his tribe Allieae because their con gested inflorescences in which the flowers are subtended by large bracts suggested the umbel-like inflorescences of Alliaceae.Species related to D. aurea but with less well-developed bracts were placed by Baker (1871) first in section Astemma of Massonia and later in subgenus Astemma of Polyxena (Baker 1897).The species recog nized by Baker (1871) in subgenus Astemma are those that are currently recognized as D. comata.D. marginata and D. zeyheri and he remarked at the time on the simi larity between them and certain species of Massonia, with the suggestion that the two groups were best com bined.This opinion was followed in recent treatments of the group (Jessop 1976; Muller-Doblies & Muller-Doblies 1997) but was not the view of Rudolf Schlechter (1924), who segregated the taxa placed by Baker in Polyxena subgenus Astemma as a distinct genus, Neobakeria, nor of Phillips (1951) or Dyer (1976).At the same time that Schlechter described Neobakeria, he also established another genus, Androsiphon, for a florally unusual species discovered a few years previously by his brother.Following this, yet another monotypic genus was established by Barker (1936), for the species Amphisiphon stylosa.These two genera were combined by Phillips (1951) but have been retained as distinct by subsequent workers, starting with Dyer (1976) and Jessop (1976).Recent studies on the molecular system atics of the group reject the recognition of these genera.The characters on which these various genera were based are best seen as extreme adaptations to a variety of polli nation and seed dispersal strategies (Goldblatt & Manning 2000).Those species with more conventional flowers that were previously placed in Neobakeria are now also confirmed to be closely allied to these florally more unusual species.Although all of these genera were included in an expanded concept of the genus Daubenya (Goldblatt & Manning 2000), not all of the species were transferred to that genus and this is accordingly done here.

SYSTEMATIC TREATMENT
Daubenya Lindl. in Botanical Register 21: t. 1813 (1835); Baker: 394 (1871); Baker: 417 (1897); E. Phillips: 193 (1951) Deciduous geophytes.Bulb turbinate to globose, sometimes deeply buried; outermost tunics leathery, dark brown, extending in a neck as narrow, flat, papery seg ments.Leaves 2, spreading to prostrate, lanceolate to ovate, longitudinally striate, glossy green, bases clasping peduncle for some distance.Inflorescence corymbose or racemose to subspicate, congested and capitate to coni cal; peduncle subterranean in flower but elongating from base in fruit and then well-exserted; bracts usually small, rarely large, usually increasing in size acropetally, the uppermost sterile, often forming a coma; pedicels suberect, vestigial to well developed in lower flowers but decreasing in length acropetally, sometimes elongating slightly in fruit.Flowers white to lilac, greenish yellow or red.strongly scented or unscented, actinomorphic or sometimes dimorphic with lower flowers slightly or strongly bilabiate, sympetalous; perianth tube cylindrical or somewhat dorsoventrally compressed, shorter in upper flowers, sometimes either adaxial or abaxial part of tube longer than opposing part in lower flowers; tepals spreading to suberect or recurved, rarely erect and con nate, linear to oblanceolate, usually subsimilar but some times lower three much enlarged.Stamens erect or slightly spreading; filaments subequal or adaxial ones slightly longer, free or fused below into short or long tube, inserted on base of tepals but sometimes tepals fused above fila ment insertion and then filaments apparently arising within perianth tube: anthers dorsifixed.Ovary ovoid; style usually between two-thirds and as long as fila ments; stigma penicillate; ovules 6-8 in two series per locule.Infructescence toppling over at maturity through elongation of lower part of peduncle.Capsule papery, obovoid, and 3-angled or sometimes somewhat inflated and 3-winged, acute or retuse at apex, usually dehiscing loculicidally from top, rarely indehiscent.style rarely persistent as a beak.Seeds globose, smooth and glossy, black.
Species eight, South Africa, mainly the more arid winter rainfall parts.Bulb subglobose, 20-35 mm diam., deeply buried; outer tunics leathery, dark brown, extending in a papery neck up to 30 mm long.Leaves 2, spreading to prostrate, ovate to lanceolate, 60-150 x 30-90 mm, bases clasping peduncle for up to 80 mm, dark green or flushed maroon.Inflorescence subspicate, capitate, exserted up to 20 mm above leaves; bracts increasing in size acropetally, lower most linear or awl-shaped, white, 7-12 mm long, those above gradually becoming broader, ovate to lanceolate, up to 10 x 2 mm, uppermost sterile, forming a coma, green or flushed pinkish, up to 8 x 3 mm; pedicels vesti gial, lowermost up to 1 mm long, flowers subsessile.Flowers white or flushed pink, strongly scented during the day and night, fragrance sweet and spicy; weakly dimorphic, lower flowers slightly bilabiate, upper flowers actinomorphic; perianth tube cylindrical or abaxial sur face flattened, lightly curved outwards, (25-)30-35(^5) mm long in lower flowers, ± 15 mm long in upper flow ers, 2-3 mm diam.; in lower flowers adaxial half of tube ± 1 mm longer than abaxial half, with short sinus between upper three and lower three tepals; tepals spreading from base and then recurved or coiling downwards, linearoblong to narrowly oblanceolate, up to 7 x 1.5-2.0mm.Stamens erect at anthesis, later spreading slightly above; filaments subequal or adaxial ones slightly longer, free, inserted on or up to 1 mm above base of tepals, (7-) 10-15 mm long; anthers pink, 2.5-3.0 mm long before dehis cence.Ovary ovoid, ± 3 mm long; style reaching to just below or slightly above filaments, 20-40 mm long.Capsule obovoid, 3-angled but rounded on angles, 10-12 x 7-8 mm.Seeds globose, ± 2 mm diam.Flowering time: mid April to mid May. Figure 1; Plate 1 A.

Key to species
Distribution and ecology: Daubenya comata is scat tered across the western portion of the South African central plateau where it is largely restricted to altitudes between 1 200-1 500 m, bordering the western rim of the drainage basin of the Orange/Gariep and Vaal Rivers (Figure 2).The white or pink flowers are strongly and sweetly scent ed throughout the day and night.They secrete small amounts of nectar and are pollinated by pollen-and nectar-collecting bees during the day (Apis mellifera and Anthophora sp.) and by small noctuid moths during the night.
Diagnosis and relationships: D. comata is readily rec ognized by the capitate inflorescence of strongly scented, white flowers with narrow, recurved or coiled tepals and free filaments.The lowermost flowers are very slightly bilabiate through a slight asymmetry in the length of the perianth tube, in the same way but to much less a degree than in D. namaquensis.The two species also resemble one another in their free filaments but they differ in inflo rescence shape, flower colour and in the form and size of the capsules.The small, subsessile capsules in D. coma ta are rounded on the sides and quite unlike the inflated and angled or winged capsules that characterize the other species in the genus.
History: D. comata was first collected on the Farm Caroluspoort, just south of the Free State border near Noupoort, on 19 March 1813 (McKay 1943) by the natu ralist and traveller William John Burchell.Fifty years were to elapse before the species was formally published by the Kew botanist J.G. Baker in 1871, with due acknowledge ment to Burchell, who had sketched and named the species in his field notebook at the time of its original col lection.It is now known to be widespread, although scat tered, across the southern African central plateau.
Conservation status: Not endangered.
Distribution and ecology: Daubenya namaquensis is known from a few collections made in the semi-arid flats east of Okiep and Springbok at an altitude of 1 000 m (Figure 2).This region, just east of Namaqualand proper and on the extreme western edge of Bushmanland, lies on the boundary between winter and summer rainfall regions and in consequence receives both erratic late summer showers as well as some rain from the occasional winter frontal system that extends inland from Namaqualand.The species responds rapidly to autumn rains and flowers in early winter before temperatures drop too low for active growth.Plants of D. namaquensis occur in small colonies in deep red sands in a vegetation dominated by the willowy shrub.Sisyndite spartea (Zygophyllaceae) and the tussock grass, Stipagrostis namaquensis.The colonies are restricted to lower-lying drainage areas in which the water table remains nearer the surface.The very deeply seated bulbs are an obvious adaptation enabling the plants to make use of this soil moisture.
The greenish yellow flowers are strongly and sweetly clove-scented throughout the day and night.They secrete small amounts of nectar and are probably pollinated by a variety of diurnal and nocturnal insects, including bees and moths.
Diagnosis and relationships: D. namaquensis is distin guished by the well-exserted conical inflorescence of strongly scented, yellow flowers with linear tepals and free filaments.Although confused by Jessop (1976) with D. marginata, the resemblance between the two species is very superficial, encompassing nothing more than the prominent inflorescence topped by a coma of sterile bracts.The flowers of D. marginata are unscented, with ovate, whitish or pale yellow tepals and golden yellow or orange filaments that are fused at the base into a short col lar.A distinct feature of D. namaquensis is the bilabiate lower flowers in which the upper side of the perianth tube is prolonged for ± 2 mm beyond the lower side, resulting in a distinct sinus between the three upper and three lower tepals.This differential in the tube length rapidly becomes less evident higher up the inflorescence and the upper flowers are quite actinomorphic.This asymmetry in the lower flowers is not very evident without careful examination, especially in pressed material.It occurs also, although to a lesser degree, in some other species of Daubenya, particularly D. comata and D. stylosa.
History: D. namaquensis was first collected by Max Schlechter during a trip to Namaqualand with his broth er, the botanist Friedrich Richard Rudolf Schlechter, soon after his arrival in South Africa in May 1896.Two decades were to pass before it was described, when it formed the basis of the genus Neobakeria, established by Rudolf Schlechter to accommodate the group of species until then treated as Polyxena subgenus Astemma.These seven species were distinguished from Polyxena s. str.by their uniseriate stamens, and from Massonia by their inconspicuous bracts.Although struck by the wellexserted, foxtail-like inflorescence of D. namaquensis, Schlechter ironically overlooked the even more diagnos tic zygomorphy of the lower flowers through dissecting only the more accessible upper flowers, which are per fectly actinomorphic.This characteristic was also over looked by Jessop (1976), who confused the species with D. marginata.The curious zygomorphy of the lower flowers of D. namaquensis was only recently highlight ed by Miiller-Doblies & Miiller-Doblies (1997).They used it as the basis for restricting the circumscription of the genus Neobakeria to this species alone, while in turn overlooking the fact that floral zygomorphy of the same type occurs, albeit to a lesser extent, in both D. comata and D. stylosa.Over a century was to pass before the species was collected for the second time and it is still known from only a handful of specimens.Bulb subglobose, 20-30 mm diam., shallowly buried; outer tunics leathery, dark brown, extending as a papery neck up to 40 mm long.Leaves 2, spreading, ovate to elliptic, 50-100 x 25-50 mm, base clasping peduncle for 20-30 mm, apiculate, dark green or flushed maroon.Inflorescence subspicate, capitate or conical, exserted 20-60 mm above leaves at flowering; bracts erect, low ermost triangular, 2-5 mm long, becoming oblanceolatespathulate acropetally, up to 8 x 3 mm, uppermost ster ile, imbricate, forming a short or elongate coma flushed orange, up to 10 x 5 mm; pedicels 1.0-2.5 mm long.Flowers greenish yellow with yellow or orange stamens, unscented; perianth tube compressed-cylindrical, 10-15(-20) x 2-3 mm; tepals suberect, inner three con nate for 0.5-1.0mm beyond outer three, lanceolate, conduplicate, 6-8 x 2.5 mm.Stamens connate for 1.5-2.5 mm in a wide-mouthed, slightly flaring staminal collar; filaments suberect and lightly incurved, 10-15(-22) mm long; anthers yellow, 2.0-2.5 mm long before anthesis.Ovary ovoid, ± 6 mm long; style lightly deflexed, 11-20 mm long.Capsule abscising at maturity, ellipsoid to obovoid, 12-15 x 15-20 mm, somewhat inflated, introrse at base, obtuse at apex, 3-angled below but 3-winged at apex.Seeds globose, ± 2.5 mm diam., glossy black.Flowering time: May to July, rarely to August.Figure 4; Plate 1C.
Distribution and ecology: Daubenya marginata is widespread across the Roggeveld Escarpment and west ern karoo at altitudes of 1 000-1 500 m, from near Calvinia and Williston southwards to Sutherland, thence eastwards along the Nuweveld scarp as far inland as Fraserburg and Loxton (Figure 2).A few isolated popu lations occur to the west, below the Bokkeveld Es carpment, at an altitude of 200 m in the Knersvlakte north of Vanrhynsdorp.Populations occur on silt or grit ty clay, more rarely red clays derived from dolerite, in seasonally moist depressions or washlines.D. marginata grows sympatrically with D. aurea in some colonies but flowers several months earlier.
Diagnosis and relationships: D. marginata is mor phologically very close to D. zeyheri and the two were regarded as conspecific by Jessop (1976).There are, however, several differences between them, some rather subtle.In D. marginata the inflorescence is typically conical and protrudes above the leaves, and is invariably surmounted by a coma of conspicuous yellow or orange, often spathulate bracts.The globular inflorescence of D. zeyheri, in contrast, does not protrude much above the leaves and is at most topped by an inconspicuous coma of narrow, green bracts.The two species differ also in flower form and colour, although in both the perianth tube varies greatly in length.In D. marginata the peri anth tube is always conspicuously flattened and 2-3 mm in diameter, the tepals tend to be flushed yellow and hrm-textured and the filaments are uniformly pigmented yellow to orange or red.The flowers of D. zeyheri, in contrast, are characterized by a much more slender peri anth tube 1.0-1.5 mm in diameter, white tepals that tend to be almost papery in texture and, most conspicuously, reddish filaments that are flushed deep purple at the base where they are fused into a shallow collar.
The plants with their orange to reddish stamens and conical inflorescence topped with enlarged orange or yellow bracts are conspicuous when in flower and are visited by Malachite sunbirds which probe the flowers for nectar.The relatively broad tubes contain abundant nectar that accumulates in the wide-mouthed staminal collar where it is readily accessible to the birds, which feed from the ground.The lack of floral scent is also con sistent with bird pollination.
History: the true identity of this species has been mis understood ever since the Kew botanist J.G. Baker con fused it with Massonia angustifolia L.f., possibly as early as 1871 but certainly by 1897.This mistake was perpetuated by subsequent authors until recently, when it was shown that M. angustifolia is actually conspecific with Massonia echinata L. (Manning & Van der Merwe 2002).The species known by all authors after Baker as M. angustifolia is in fact properly known as Daubenya marginata, first described as Massonia marginata by the Berlin Botanist Carl Kunth.Although known to his pre decessor, Carl Willdenow, it was not formally named until Kunth (1843) published the name that had earlier been suggested for it by Willdenow.The collector and place of collection of the type specimen are unknown.The species was later described as Polyxena haeman thoides by Baker in 1888, based on a plant collected by the Cape Town stockbroker and botanist, Harry Bolus, on the Nuweveld Mountains near Fraserburg in April 1886.It seems that Bolus intended describing the species himself, as Haemantholirion capense, but Baker must have dissuaded him from this step.Baker did, however, retain the reference to the genus Haemanthus (Amaryllidaceae) that the appearance of the plant had suggest ed to Bolus.Baker, although suspecting that his species was similar to Kunth's, had not seen material of M. mar ginata and thus described it as new.The species was re cognized until Jessop (1976) revised the genus.Un convinced that the apparent differences between it and D. zeyheri were significant, he combined the two taxa under the misapplied name M. angustifolia.Conservation status: Not endangered.Bulb subglobose, 20-30 mm diam., outer tunics leath ery.dark brown, extending as a papery neck up to 10 mm long.Leaves 2, spreading, elliptic to lanceolate, 60-150 x (10-)30-80 mm.bases clasping peduncle for 10-100 mm.apiculate, dark green.Inflorescence corymbose, capitate, exserted 20-40 mm above leaves; bracts erect, lowermost triangular.2-3 mm long, becoming lanceolate acropetally and increasing in size, up to 8 x 2 mm.uppermost sometimes sterile and forming inconspicuous coma: pedicels 2-10 mm long, lowermost sometimes longer than upper.Flowers translucent, whitish tinged pink on tube, stamens reddish tinged purple at the base, papery in texture, unscented: perianth tube cylindrical, lightly constricted at mouth.12-20 mm long in lower flowers but rarely more than 13 mm in upper flowers, 1.5-2.0mm diam.inner tepals connate for 0.5-1.0mm beyond outer ones; tepals spreading at base then suberect, lanceolate, conduplicate, 6-10 x 2.5-3.0 mm.Stamens reddish but staminal collar purple, connate for 1.0-1.5 mm in a flaring staminal collar; filaments suberect, 12-19 mm long; anthers yellow or reddish, 2-3 mm long before anthesis.Ovary ovoid.± 4 mm long; style 15-30 mm long.Capsule abscising at maturity, broadly ovoid, 1 0 -1 5 x 9 -1 3 mm, somewhat inflated, 3angled, introrse at base, obtuse at apex.Seeds globose, ± 2.5 mm diam., glossy black.Flowering time: late May and June to early July.Figure 5; Plate ID.
Distribution and ecology: Daubenya zeyheri is a coastal species occuring near sea level, unlike the other species in the genus, which occur at higher altitudes inland.It is restricted to the west coast of Western Cape, where it is now known from only three locations, one at Paternoster and Cape Columbine and the others less than 20 km to the south, near Saldanha (Figure 2).The material that formed the type of M. pedunculata was collected 10 km southeast of Hopefield on the Farm Schaapplaatsfontein but no fur ther collections have been made from here and this popu lation has presumably disappeared under wheat.D. zeyheri grows on sandy calcareous soils overlying limestone, sometimes in dense colonies.
The flowers, with their reddish filaments and con trasting glistening purple centres attract the attention of Lesser double-collared sunbirds, which probe the flow ers for the copious nectar that is held in the wells formed by the staminal collars.The lack of floral fragrance is typical of bird-pollinated flowers.
Diagnosis and relationships: although closely related to D. marginata and sometimes difficult to separate from it in the dry state, living plants of D. zeyheri are readily recognized by the conspicuous purple centres to the flowers.The pigmentation is restricted to the staminal collar and when this is filled with nectar the flowers appear to have a glistening black centre that is quite lack ing in D. marginata, in which the stamens are uniformly yellow or orange to reddish.Other more subtle differ ences between the two species are discussed under D. marginata.The two were treated as conspecific by Jessop (1976).
History: D. zeyheri has been described several times under different names but the earliest name dates from 1843, when the species was described by the Berlin botanist, Carl Kunth, from a collection made by the pro fessional plant collector, Carl Zeyher.Zeyher's collec tion was made sometime between 1829 and 1834, prob ably near Saldanha Bay, which is still one of only two sites where the species is known to occur.The species was redescribed twice, both times by the Kew botanist J.G. Baker.In 1892 he described Massonia pedunculata from a specimen collected near Hopefield by the German naturalist Frans Bachmann, who practised medicine in the village from 1886-1887.A few years later he named Massonia burchellii from a collection made by the natu ralist and traveller William John Burchell.Baker was clearly unsure about the exact identity of several of his species because he later transferred M. burchellii to Polyxena while leaving M. pedunculata in Massonia, although noting that it lacked the large bracts that are so characteristic of the genus.

D aubenya alba
Distribution and ecology: Daubenya alba is known from a few scattered localities along the edge of the Roggeveld Escarpment between Calvinia and Middelpos at an altitude of 1 000-1 500 m (Figure 7).The plants
Diagnosis and relationships: D. alba is readily recog nized by its white to lilac flowers with the stamens fused below into a narrow tube 6-10 mm long.The slight zygomorphy of the lowermost flowers, resulting from a slight asymmetry in the staminal tube, that is mentioned in the original description, is not always evident.The relationships of D. alba appear to lie with D. capensis, which it resembles in several respects, particularly its relatively long pedicels, cylindrical staminal tube and large, obtriangular capsules that are deeply retuse at the apex.The two species are readily distinguished by their flowers.In D. capensis the firm-textured flowers are bright yellow with a shorter, broader tube, 6-10 mm long and ovate tepals.The flowers of D. alba are altogether more delicate, with a slender perianth tube, 12-20 mm long and narrowly oblanceolate tepals.In addition D. alba lacks the conspicuous disc occluding the mouth of the staminal tube that is characteristic of D. capensis.
History: the most recently discovered species in the genus, D. alba was in cultivation in a few specialist col lections by the early 1990s, although the type material was only collected from the wild in June 2001.
Distribution and ecology: Daubenya capensis is endemic to the immediate vicinity of Nieuwoudtville, where it is known from several populations around the village at an altitude of 800 m (Figure 7).It is restricted to seasonally moist dolerite flats in red clay.The large, prominently three-winged capsules borne on relatively long pedicels form a rounded, balloon-like infructes cence that is readily dispersed by the wind.
The curious disc that occludes the mouth of the tube is glossy and has the appearance of being wet even when dry, although nectar does in fact ooze out of the pore in the staminal disc and accumulate on it.Nothing is known about the pollination biology of the species but the firm texture of the flowers and their yeasty odour suggest rodent pollination.
Diagnosis and relationships: D. capensis is readily distinguished from other species of Daubenya in which the filaments are fused into a long, narrow column by the peculiar disc that occludes the mouth of the column and through which the style protrudes via a narrow pore.D. capensis is most similar in floral form to D. alba, which it also resembles in its distinctly pedicellate flowers and large, obtriangular capsules that are deeply retuse at the apex.The fruits in these two species are among the largest in the genus.D. capensis is readily distinguished from D. alba by its more robust, yellow flowers with shorter perianth tube, 6-10 mm long and by the conspicu ous disc that occludes the mouth of the staminal tube.
History: D. capensis was first collected in July 1897 by the German botanist, Friedrich Richard Rudolf Schlechter and was described by him some years later, when it formed the basis of his monotypic genus Androsiphon.
Here it remained until recently, when that genus was included in Daubenya (Goldblatt & Manning 2000).
Distribution and ecology: Daubenya capensis is endemic to the immediate vicinity of Nieuwoudtville, where it is known from several populations around the village at an altitude of 800 m (Figure 7).It is restricted to seasonally moist dolerite flats in red clay.The large, prominently three-winged capsules borne on relatively long pedicels form a rounded, balloon-like infructescence that is readily dispersed by the wind.
The curious disc that occludes the mouth of the tube is glossy and has the appearance of being wet even when dry, although nectar does in fact ooze out of the pore in the staminal disc and accumulate on it.Nothing is known about the pollination biology of the species but the firm texture of the flowers and their yeasty odour suggest rodent pollination.the filaments are fused into a long, narrow column by the peculiar disc that occludes the mouth of the column and through which the style protrudes via a narrow pore.D. capensis is most similar in floral form to D. alba, which it also resembles in its distinctly pedicellate flowers and large, obtriangular capsules that are deeply retuse at the apex.The fruits in these two species are among the largest in the genus.D. capensis is readily distinguished from D. alba by its more robust, yellow flowers with shorter perianth tube, 6-10 mm long and by the conspicu ous disc that occludes the mouth of the staminal tube.
History: D. capensis was first collected in July 1897 by the German botanist, Friedrich Richard Rudolf Schlechter and was described by him some years later, when it formed the basis of his monotypic genus Androsiphon.
Here it remained until recently, when that genus was included in Daubenya (Goldblatt & Manning 2000).

7.
Daubenya stylosa (W.F.Barker) A.M.van der Merwe & J.C. Manning, in Goldblatt & J.C.Manning in Strelitzia 9: 713 (2000).Bulb subglobose to turbinate, 20-35 mm diam., usu ally shallowly buried; outer tunics leathery, dark brown, extending in a papery neck up to 5 mm long.Leaves 2, suberect to spreading, ovate to lanceolate, (20-)50-100 x 15-50 mm, bases clasping peduncle for up to 30(-60) mm, dark green.Inflorescence subspicate, capitate or rarely conical, exserted up to 50 mm above leaves; bracts increasing in size acropetally, lowermost lanceolate, white, 3-5 mm long, uppermost oblanceolate, up to 8 x 3 mm; pedicels suberect, 1-3 mm long, flowers subsessile.Flowers greenish yellow with free part of filaments and style golden yellow, honey-scented; perianth tube cylindrical, 8-14 mm long in lower flowers with portion below level of filament insertion 4-7 mm long but 6-8 mm long in upper flowers with portion below level of fil ament insertion 2-3 mm long, 2.0-2.5 mm diam.; lower flowers lightly bilabiate with adaxial portion of tube ± 3 mm longer than abaxial half through a greater degree of fusion of upper three tepals; tepals connate into tube above level of filament insertion, free parts oblong to ovate, 2-5 mm x 2.0-2.5 mm.Stamens fused below into tube 10-15 mm long, free parts of filaments suberect, 9-10 mm long; anthers yellow, up to 2.5 mm long before dehiscence.Ovary ovoid, ± 3 mm long; style reaching to top or just beyond filaments, 17-28 mm long.Capsule depressed-ovoid, 10-12 x 7-10 mm, lightly retuse at base and tapering above into peristent style which forms slender beak ± 20 mm long, indehiscent but eventually separating along septa.Seeds globose, ± 2 mm diam., glossy black.Flowering time: mid May to mid June. Figure 9; Plate 1G.Distribution and ecology: Daubenya stylosa is a high ly local endemic around Nieuwoudtville where it is known from a few populations in the immediate vicinity of the town, especially along the trekpath that passes the Farm Glenlyon, at an altitude of 800 m (Figure 10).The populations here are extremely dense.The plants typi cally grow in lower-lying drainage areas in red doleritic clays.Scattered plants occur along the foot of dolerite outcrops on the Wildflower Reserve and on Glenlyon itself.The species is more or less sympatric with D. capensis but the two cannot be confused, either in flower or fruit.The compact infructescence in D. stylosa lacks the adaptations for wind dispersal typically found in most other species in the genus and the seeds are mainly shed immediately around the parent plants.
The sweetly scented, greenish yellow flowers secrete small amounts of nectar that ooze out of the mouth of the staminal column.They are visited avidly for the nectar by honeybees (Apis mellifera) and the Painted lady but terfly (Cynthia cardui) during the day and by several species of noctuid moth at dusk.Diagnosis and relationships: the species is readily rec ognized by its almost completely fused tepals, which form a tube around the staminal column, and by the persistent styles that remain attached to the capsule.These stiff, prong-like beaks are a conspicuous feature of the subsessile fruits.In addition, unlike most other species in the genus, which have loculicidally dehiscent capsules, the fruits in D. stylosa are essentially indehiscent although the locules eventually separate along the septa to release the seeds.History: D. stylosa was first collected relatively recently, in June 1934, by retired Royal Navy Paymaster, T.M. Salter, best known for his work on the genus Oxalis.It was described soon thereafter in the monotypic genus Amphisiphon by the Cape Town botanist W.F. Barker.Included in Amphisiphon by Phillips (1951), the genus was recently transferred to Daubenya (Goldblatt & Manning 2000).Bulb subglobose to turbinate, 20-35 mm diam., usu ally shallowly buried; outer tunics leathery, dark brown, extending in a short, papery neck up to 10 mm long.Leaves 2, suberect to spreading, ovate to lanceolate, 50-100(-140) x 20-75 mm, bases clasping peduncle for 5-50 mm, dark green.Inflorescence corymbose, capitate, exserted up to 50 mm above leaves; bracts decreasing in size acropetally, greenish, lowermost obovate to oblance olate, 25-35 x 10-15(-20) mm long, uppermost up to 20 x 10 mm; pedicels suberect, lowermost 5-10 long, uppermost up to 3 mm long.Flowers brilliant red or canary yellow, rarely flushed reddish at tips of tepals, at most lightly scented, ± zygomorphic and bilabiate; lower flowers strongly bilabiate with abaxial or lower half of tube much prolonged and adnate to filament column and lower three tepals forming conspicuous lip separated from upper tepals by pronounced sinus, sometimes dor sal or uppermost tepal arising very much below remain ing tepals, perianth tube obliquely cylindrical, 15-25 mm long on upper side but 20-40 mm long on lower side, 3 ^ mm diam., upper tepals oblong-ovate, laterals obliquely so, 2-4 x 1.5-2.0mm, lower tepals oblanceo late to obovate, subequal or median larger, 20-40 x 5-15 mm; upper flowers weakly zygomorphic or actinomor phic, perianth tube 13-20 x 2.5-3.5 mm, tepals oblong, 2 -\ x 1.5-2 mm.Stamens fused basally into an oblique tube, 5-7 mm long on adaxial side but much longer on abaxial side, especially in lower flowers, and there adnate to lower tepals such that lower stamens appear to be free, free parts of filaments suberect, 4-8 mm long; anthers yellow, 2.5-3.0 mm long before dehiscence.Ovary ovoid, 7-8 mm long; style reaching to top or just beyond all or only adaxial filaments, 20-25 mm long.Capsule oblong-ovoid, 3-angled, 15-20 x 7-10 mm, introrse at base but acute at apex.Seeds globose, ± 3 mm diam., glossy black.Flowering time: late August to mid September.Figure 11; Plate 1H.
Distribution and ecology: D. aurea is a highly local ized endemic of the Roggeveld Escarpment known from three or four localities midway between Sutherland and Middelpos, at an altitude of ± 1 500 m (Figure 10).The known populations lie along the eastern foot of a mean dering dolerite ridge, with the total distribution little more than 10 km in extent.The plants grow in colonies in low-lying drainage lines in seasonally damp red clay derived from dolerite.Populations are sympatric with D. marginata in several of the localities but flower much later than that species.D. aurea occurs in two colour forms, the more common one a brilliant red and the other canary yellow.Populations are typically uniformly coloured, although red populations will often contain occasional yellow morphs, whereas yellow populations may include a few plants in which the tepals are flushed reddish at the tips.The yellow populations consistently come into flower one or two weeks before the red but the fruits appear to take much longer to ripen.The prominent papery bracts that are characteristic of the species are particularly conspicuous in fruit, serving as sails that assist in the wind dispersal of the infructescences.
The inflorescence resembles a large daisy capitulum in the genus Arctotis or Gazania due to the enlargement of the lower florets, and the species is adapted to polli nation by monkey beetles (Scarabidae: Holpiinae) that congregate on the large flower heads to mate.The flow ers, unlike those of the other species in the genus, do not produce nectar and are also unscented or at most weakly scented.Both of these characteristics are typical adapta tions to pollination by monkey beetles.
Diagnosis and relationships: D. aurea is a highly dis tinctive species that cannot be confused with any other species in the genus.The highly zygomorphic lower flowers in which the outer tepals are greatly exaggerated in size make this species instantly recognizable.It is also characterized by large floral bracts that are persistent and papery in fruit.
History: it is ironic that the source of the original material of the best known species of Daubenya remained obscure for so long.D. aurea was described in 1835 by John Lindley, Professor of Botany at the University College, London and head of the Chelsea Physic Garden, from a yellow-flowered plant cultivated by the nurserymen.Messrs Young of Epsom.Nothing is known of its origin beyond that the Youngs obtained their plant from the Cape of Good Hope under the name Massonia lutea.Three years later a reddish-flowered variant of the species came to Lindley's attention through the agency of John Royle.then Professor of Materia Medica at King's College.London, from the collection of Robert Barchard of East Hill.Wandsworth.Lindley was struck by the apparently more extreme zygomorphy evident in the flowers of the red form and he recognized this collection as a different species, D. fulva.on this basis.The name is an allusion to the tawny colour of the plant, but this, along with the rather etiolated appearance of the plant, was probably the result of cultivation in less than ideal light.From the illustration it is clear that this plant was in fresher flower than the first one that Lindley had seen and he presumably examined only the lowest, most zygomorphic flowers in the inflorescence.This plant had been received from the Cape of Good Hope with other bulbs but was believed to have been collected somewhere along the East Coast of Africa or in Madagascar.This was, of course, not the case but it is possible that Barchard received his plant from the same source as did the Youngs.A dried plant of D. fulva was later seen bv Baker (1897) in the herbarium of William Wilson Saunders, who had a notable garden at East Hill from 1857 to 1874 and was a neighbour of Barchard.It is thus probable that his specimen came from Barchard himself and is possibly even the one illustrated in the protologue.At the same time that Lindley was describing his two species of Daubenya, a specimen appears to have been in the possession of William Henry Han ey in Cape Town.Harvey, although nominally Colonial Treasurer, occupied most of the four years that he spent in Cape Town between 1835 and 1842 on botany.Harvey's speci men formed the basis of D. coccinea, described by Baker (1871), which was distinguished from D. fulva by its compact inflorescence and more robust flowers.Although the provenance of the plant is unknown, its appearance is typical of wild collected material or that grown under good light, such as would have been expe rienced in Cape Town.It is impossible to be certain on this point but the more or less coincidental appearance of material in three different hands strongly suggests that it all originated from the same source.Circumstantial evi dence favours Capt Walter Synnot as this source.Synnot spent four years in Clanwilliam as deputy landdrost.returning to Ireland in 1825 with numerous bulbs and seeds.These were disposed of in England to various nurseries and included various species from the Bokkeveld and Roggeveld Escarpments, some of which only flowered in the 1830s.It is thus quite possible that among them were seeds or plants of D. aurea.
Almost a century was to pass without further infor mation until, in 1920.Rudolph Marloth came upon Daubenya aurea in the wild on the Roggeveld Escarpment.Marloth.a retired chemist with a passionate interest in South African botany, sent plants and exact details of the locality and habitat to the Pretoria botanist.E.P Phillips, who published the story of the rediscovery in 1922.At the same time it became evident that no significant structural differences existed between the three species hitherto known and they were consequently reduced to one.Although further plants were sent to Kirstenbosch gardens by a Mr Metelerkamp in 1936 (Hall 1970), they soon languished and it was left to Harry Hall, then Senior Horticulturalist at the Gardens, to locate Marloth's collecting site in 1968 and bring the species into cultivation once more.He has provided a lively account of his search, illustrated with fine photo graphs of the plants in the wild (Hall 1970).
Diagnosis and relationships: D. capensis is readily distinguished from other species of Daubenya in which
It has been recorded from Beaufort West near the northern border of the Western Cape, eastwards to Molteno in the Eastern Cape, through the Free State and eastern parts of the North-West as far north as Barberspan near Delareyville.Plants grow in colonies in seasonally waterlogged loam or clay, particularly the red, glutinous clays derived from the doleritic sills that are frequently exposed across the interior plateau.These soils retain moisture and remain damp for longer than the surrounding karoo shales.Like D. namaquensis, another early flowering species, D. comata responds to late sum mer showers, flowering before the soils dry up and the temperatures drop too low for active growth.It shares this strategy and habitat with several other sympatric dwarf geophytes with a similar growth habit, particular ly Androcymbium asteroides, Moraea falcifolia and Polyxena ensifolia.