A taxonomic revision of the southern African native and naturalized species of Silene L . ( Caryophyllaceae )

The native and naturalized species of Silene L. in southern Africa are reviewed, with full synonomy and the description of two new species from the West Coast of Western Cape. Eight native species and three naturalized species are recognized, including the first identification in southern Africa of the Mediterranean S. nocturna L. The identity of S. aethiopica Burm., which has remained unknown since its description, is established and is found to be the oldest name for S. clandestina Jacq. Patterns of morphological variation within each species are discussed and subspecies are recognized for geographically segregated groups of populations that are ± morphologically diagnosable. The following new names or combinations are made among the southern African taxa: S. aethiopica subsp. longiflora; S. burchellii subsp. modesta, subsp. multiflora, and subsp. pilosellifolia; S. crassifolia subsp. primuliflora; S. saldanhensis; S. rigens; and S. undulata subsp. polyantha. Each taxon is described, with information on ecology and distribution, and most species are illustrated, including SEM micrographs of the seeds. * Compton Herbarium, South African national Biodiversity Institute, Private Bag X7, Claremont 7735, South Africa / Research Centre for Plant growth and Development, School of life Sciences, university of KwaZulu-natal, Pietermaritzburg, Private Bag X01, Scottsville 3209, South Africa. E-mail: J.Manning@sanbi.org.za. ** B.A. Krukoff Curator of African Botany, Missouri Botanical garden, P.O. Box 299, St. Louis, Missouri 63166, uSA. E-mail: peter.goldblatt@mobot.org. MS. received: 2011-12-19. 148 Bothalia 42,2 (2012) exist for parts of north Africa (Wickens 1976; gilbert 2000) and much of sub-Saharan Africa (Turrill 1956a; Wild 1961) but the taxonomy of Silene in southern Africa still rests largely on Sonder’s (1860) treatment in Flora capensis, and only minor advances in our understanding of the southern African species have been made since then (Rohrbach 1869, 1869–70; Bocquet 1977; Masson 1989; goldblatt & Manning 2000). The South African species of Silene are unique among the flora of the subregion in having been fundamentally misunderstood by Thunberg (1794), who misidentified the majority of his collections as European species (Table 1), leaving it to later authors to recognize that they were indigenous to the country and undescribed. Burman (1768) had earlier made the same mistake, identifying as European all but two (S. crassifolia L. and S. aethiopica Burm.) of the eight species that he listed from the Cape. Two previous attempts at revising the southern African species, by g. Bocquet in the 1970s and D. Masson in the 1980s, were never completed but many of the southern African herbarium collections bear annotation labels reflecting Masson’s unpublished species concepts and names, adding further to the confusion. This revision of Silene in southern Africa aims to redress the situation and provide a base for future studies of the genus in the subcontinent. This is especially important as the South African winter rainfall region is a minor centre of diversification for Silene in Africa. Species delimitation in the genus is notoriously difficult and most African species, especially those with a wider distribution, display a complex and often confusing pattern of local variation that defies formal classification (gilbert 2000). The primary reason for this is almost certainly that the species are often facultatively autogamous, which facilitates the differentiation of local phenotypes, probably independently and repeatedly. Population genetic studies are likely to shed significant light on the nature and origin of these local forms. We have recognized as species only entities that are morphologically (and preferably also ecologically) discontinuous, based on the assumption of ± complete interspecific infertility. Although Stace (1978) observed that interspecific hybridization is generally rare in Silene, recent molecular investigations (e.g. Erixon & Oxelman (2008) and Frajman et al. (2009)) suggest that it is an important consideration in some species groups. Where possible, we recognize infraspecific taxa for geographically coherent groups of populations that show evident but incomplete morphological differentiation. We have uniformly applied the rank of subspecies to these taxa but experimental investigation might indicate that other ranks may be useful in some instances, such as in S. aethiopica, where subsp. longiflora may be better treated at the level of forma. The genetics of the maritime forms of S. aethiopica and S. undulata also require investigation. Our intention, therefore, is to present a preliminary classification that can form the basis for future experimental studies in the genus. Another intriguing aspect of the genus is its history and diversification in southern Africa. The indigenous species are currently distributed among three sections, all of which are best represented in the northern hemisphere and presumably originated there. Silene burchellii (sect. Fruticulosae) is widely distributed through the eastern half of the continent, from Ethiopia (and Arabia) to the Cape Peninsula in the extreme south, and the most parsimonious interpretation is that the species migrated from north Africa southwards along the East African Afromontane corridor. Silene undulata (sect. Elisanthe), in contrast, is largely southern African, extending only as far north as the highlands of eastern Zimbabwe, and its origin in southern Africa remains unclear but long-distance dispersal appears to be more likely. Both sections have subsequently undergone minor adaptive radiation in the coastal regions of the subcontinent. The third lineage is represented by S. aethiopica (sect. Dipterospermae), the only native southern African annual species. Essentially endemic to the winter-rainfall region in the extreme south and southwest, with no track in tropical Africa, S. aethiopica most likely arrived in the subcontinent through long-distance dispersal. MATERIALS AnD METHODS All relevant types were examined, as well as all herbarium material from BOL, MO, nBg, PRE, and SAM (acronyms after Holmgren et al. 1990), the primary collections of species from southern Africa. All species were also studied in the field. Ripe seeds from dehisced capsules were mounted directly on aluminium stubs and coated with gold-palladium before viewing at the SEM unit, university of Cape Town. Seeds were sampled from at least two different collections per species (Figures 1, 2).


InTRODuCTIOn
Silene L. (tribe Sileneae), with 600-700 species (greuter 1995b), is distributed mainly through the temperate regions of the northern Hemisphere, with its principal centre of diversity in the Mediterranean and Middle East (Oxelman & Lidén 1995).Over 90 species are recorded in Africa, the vast majority in north Africa, with very few extending southwards into sub-Saharan Africa.Just three native and one introduced species of Silene are known from south tropical Africa (Turrill 1956a;Wild 1961), with several additional species currently accepted in southern Africa.
Consensus has still to be reached on the delimitation of Silene.The circumscription of the genus had been gradually expanded over the last few decades of the twentieth century until it coincided essentially with the delimitation of tribe Sileneae (greuter 1995b).This consolidation was supported by a preliminary molecular sequence analysis of the tribe (Oxelman & Lidén 1995), which nested all previously recognized genera of Sileneae in Silene, with the possible exception of Agrostemma L., but had been presaged by the breakdown of morphological differences between recognized segregates when these were examined on a global scale.Cucubalus L., which is deeply nested in Silene, is distinguished primarily by its berry-like fruits, which evidently represent a specialized dispersal strategy.Lychnis L., which forms a discrete clade within Sileneae, was traditionally diagnosed by having capsules with as many teeth as styles (vs.twice as many in Silene) (e.g.Chowdhuri (1957) and Chater & Walters (1964)) but the two genera were diagnosed only with great difficulty by Bit-trich (1993) using a combination of trivial characters, and his recognition of the two appears to have been provisional until Silene could be conserved against Lychnis, under which name it had earlier been included by Scopoli (1771).The generic name Lychnis therefore took priority until it was formally rejected in favour of Silene (Brummit 1994).
There are currently two opposing views on generic delimitation in Sileneae.The first, advocated by greuter (1995b), is an inclusive one that recognizes just the two genera Agrostemma and Silene.In this sense, Silene is broadly distinguished within subfamily Caryophylloideae (Bittrich 1993) by its actinomorphic flowers with 3 or 5 styles.The alternative view, recently developed by Oxelman et al. (2000), proposes the partitioning of Silene among seven genera, of which only Lychnis (30 spp.) includes more than a handful of species.The clade that constitutes Silene in this narrow sense is only weakly supported in the molecular analysis and is also not easily diagnosed morpholgically from its segregates.Although we favour a more inclusive circumscription of the genus, the native and introduced southern African species are all included in Silene either way.
We follow the infrageneric classification of Silene s. lat.proposed by greuter (1995b), based largely on Chowdhuri (1957) with some rationalization and several nomenclatural corrections.Although there is mounting evidence (Petri & Oxelman 2011) that at least some of these sections are not monophyletic, no satisfactory alternative has yet been offered.Significant characters at sectional level are the type of inflorescence, presence or absence of septa in the ovary, number of styles, shape and venation of the calyx, number of teeth in the dehiscing capsule, and the shape of the seeds.
Four global treatments have laid the foundations for our present understanding of the genus (Otth 1824;Rohrbach 1869Rohrbach , 1869Rohrbach -1870;;Williams 1896;Chowdhuri 1957), and several modern regional treatments of the European species have also been published (greuter 1995b and references therein).Regional floristic studies Bothalia 42,2: 147-186 (2012) A taxonomic revision of the southern African native and naturalized species of Silene L. (Caryophyllaceae) Bothalia 42,2 (2012) exist for parts of north Africa (Wickens 1976;gilbert 2000) and much of sub-Saharan Africa (Turrill 1956a;Wild 1961) but the taxonomy of Silene in southern Africa still rests largely on Sonder's (1860) treatment in Flora capensis, and only minor advances in our understanding of the southern African species have been made since then (Rohrbach 1869(Rohrbach , 1869-70;-70;Bocquet 1977;Masson 1989;goldblatt & Manning 2000).The South African species of Silene are unique among the flora of the subregion in having been fundamentally misunderstood by Thunberg (1794), who misidentified the majority of his collections as European species (Table 1), leaving it to later authors to recognize that they were indigenous to the country and undescribed.Burman (1768) had earlier made the same mistake, identifying as European all but two (S.crassifolia L. and S. aethiopica Burm.) of the eight species that he listed from the Cape.Two previous attempts at revising the southern African species, by g.Bocquet in the 1970s and D. Masson in the 1980s, were never completed but many of the southern African herbarium collections bear annotation labels reflecting Masson's unpublished species concepts and names, adding further to the confusion.This revision of Silene in southern Africa aims to redress the situation and provide a base for future studies of the genus in the subcontinent.This is especially important as the South African winter rainfall region is a minor centre of diversification for Silene in Africa.
Species delimitation in the genus is notoriously difficult and most African species, especially those with a wider distribution, display a complex and often confusing pattern of local variation that defies formal classification (gilbert 2000).The primary reason for this is almost certainly that the species are often facultatively autogamous, which facilitates the differentiation of local phenotypes, probably independently and repeatedly.Population genetic studies are likely to shed significant light on the nature and origin of these local forms.We have recognized as species only entities that are morphologically (and preferably also ecologically) discontinuous, based on the assumption of ± complete interspecific infertility.Although Stace (1978) observed that interspecific hybridization is generally rare in Silene, recent molecular investigations (e.g.Erixon & Oxelman (2008) and Frajman et al. (2009)) suggest that it is an important consideration in some species groups.Where possible, we recognize infraspecific taxa for geographically coherent groups of populations that show evident but incomplete morphological differentiation.We have uniformly applied the rank of subspecies to these taxa but experimental investigation might indicate that other ranks may be useful in some instances, such as in S. aethiopica, where subsp.longiflora may be better treated at the level of forma.The genetics of the maritime forms of S. aethiopica and S. undulata also require investigation.Our intention, therefore, is to present a preliminary classification that can form the basis for future experimental studies in the genus.
Another intriguing aspect of the genus is its history and diversification in southern Africa.The indigenous species are currently distributed among three sections, all of which are best represented in the northern hemisphere and presumably originated there.Silene burchellii (sect.Fruticulosae) is widely distributed through the eastern half of the continent, from Ethiopia (and Arabia) to the Cape Peninsula in the extreme south, and the most parsimonious interpretation is that the species migrated from north Africa southwards along the East African Afromontane corridor.Silene undulata (sect.Elisanthe), in contrast, is largely southern African, extending only as far north as the highlands of eastern Zimbabwe, and its origin in southern Africa remains unclear but long-distance dispersal appears to be more likely.Both sections have subsequently undergone minor adaptive radiation in the coastal regions of the subcontinent.The third lineage is represented by S. aethiopica (sect.Dipterospermae), the only native southern African annual species.Essentially endemic to the winter-rainfall region in the extreme south and southwest, with no track in tropical Africa, S. aethiopica most likely arrived in the subcontinent through long-distance dispersal.

MATERIALS AnD METHODS
All relevant types were examined, as well as all herbarium material from BOL, MO, nBg, PRE, and SAM (acronyms after Holmgren et al. 1990), the primary collections of species from southern Africa.All species were also studied in the field.

Vernacular name: bladder campion.
Distribution and ecology: native to Europe, where it is widely distributed across the continent through the Middle East into temperate Asia; introduced into Ethiopia, Australia, South Africa (where it is cultivated as an ornamental) and also north America, where it is considered a weed.
Diagnosis: readily distinguished by the complete lack of vestiture and by the pale greenish or whitish, inflated, bladder-like calyx with 20 equal longitudinal veins joined by extensive reticulated veining throughout.The petals, deeply divided to the base of the limb with the claw abruptly narrrowed and filiform in the basal half, are also diagnostic.Flowers are either bisexual, with long, well-exserted filaments ± 18 mm long, or functionally female, with relatively short filaments, 5-6 mm long, deeply included within the calyx.
Distribution and ecology: widely distributed through the Mediterranean and long established in southwest Britain, whence it was presumably introduced to South Africa, probably as an ornamental.In South Africa the species has been recorded from mesic situations in the Western Cape winter-rainfall region (Tygerberg Hills, around Stellenbosch and Oudtshoorn, and in the Langkloof) and from the Hogsback in Eastern Cape, as a garden escape in waste places, along roadsides, around old lands, and in vineyards.It was first recorded as an escape around Stellenbosch in the early twentienth century (Figure 5).
Distribution and ecology: widely distributed through the more temperate parts of southwestern and eastern southern Africa northwards into eastern Zimbabwe (Figure 8), occurring in a variety of stony or rocky habitats, in coastal and inland scrub, often in ravines, along riverbanks and forest margins, and in moist grassland, from just above the high tide mark to well over 2 000 m.The species is absent from the more arid regions, notably Bushmanland and the Kalahari in northern Cape and the great Karoo in Western and Eastern Cape.
growth and flowering of the species in vegetated communities such as thicket margins is stimulated by clearance of the overburden by fire.The flowers are strictly nocturnal and strongly scented at night but although they may remain open for several hours on overcast mornings, sometimes until midday, they are then often no longer scented.The species is clearly pal-atable, being extensively browsed by both wild animals and livestock, but is also used traditionally as a medicine in many diseases, particularly fevers and delirium (Watt & Breyer-Brandwijk 1932).
Diagnosis and relationships: as circumscribed here, Silene undulata is a highly variable species with numerous local forms that have in common the development of relatively long, eglandular hairs, 1-2 mm long, on the lower portion of the stem, giving the base of the stems a characteristic, shaggy appearance (Figure 7C).In some instances the eglandular hairs are mixed with scattered, short, gland-tipped hairs, which typically predominate in the upper parts of the stems (Figure 7B), which are sometimes entirely glandular-haired.The vestiture of the calyx is similarly variable, usually including a mix of longer, eglandular hairs and short, gland-tipped hairs, but sometimes either entirely eglandular or entirely glandular.The lobes are, however, invariably fringed with eglandular hairs.Other sub-Saharan species in the section have a ± entirely glandular pubescent calyx, including the margins of the lobes.Another characteristic of the species is the relatively long, narrowly triangular to   7D).
The species varies also in the length and exsertion of the stamens and styles.Style length is dependent on the age of the flowers, and the styles are always included at anthesis.In many populations, however, the styles elongate markedly after anthesis, becoming well exserted by the second or third day.
Silene undulata was described from cultivated plants grown at Kew from seeds collected in the Western Cape by Francis Masson, who visited the region from 1772 to 1775 and again from 1786 to 1795.The earliest collection of the species, however, appears to be an unlocalised specimen at the British Museum collected in 1772 by Franz Oldenburg (Oldenburg 1260).Oldenburg, who visited the Cape in1771-1773, accompanied both Masson and Carl Thunberg on collecting trips (glen & germishuizen 2010).His fragmentary specimen at the British Museum (BM001010634) was initially ignored, then mistakenly identified as the Sicilian species Silene calycina C.Presl., before finally being correctly determined as S. undulata.It represents the form with a proportionally longer carpophore that was initially recognized at species level under the name S. capensis Otth.An equally fragmentary collection of the same form made by Thunberg, possibly from the same population that yielded seeds to Masson since the two men certainly collected together on the Cape Peninsula and along the West Coast where S. undulata occurs, was mistaken by him for the Mediterranean S. pendula (Table 1).Silene undulata exhibits numerous local forms, varying in habit, degree and nature of vestiture, length of calyx and carpophore, length and shape of calyx lobes, size of petals, degree of exsertion of stamens and styles, and shape and size of the capsule.Some of these have formed the basis of separate species but we are unable to identify any of these character states or combination of states that is consistent with the recognition of more than a single, variable species.The relative length of carpophore and capsule, in particular, has been used to distinguish several species in the complex, viz.S. bellidioides (carpophore one quarter as long as capsule), S. undulata (carpophore one third as long as capsule), and S. capensis (carpophore at least half as long as capsule) (Sonder 1860).A further two species were recognized on the basis of their relatively hairless leaves, S. eckloniana from Cape Recife with eglandular stems, and leaves glabrous above but hairy on the margins and along the midrib beneath, and S. caffra, also from Eastern Cape, with subglabrous stems and leaves that are ciliolate along the margins but otherwise glabrous on both surfaces.With numerous collections now to hand, we are unable to correlate any of these differences with other character states, nor with geography, and conclude that they represent independent and local variants within a single species.
Some of these names were consolidated by Rohrbach (1869), who treated S. caffra, S. capensis, and S. diurniflora as synonyms of S. undulata.Of the remaining names, only S. bellidioides and S. undulata have been applied in recent years, always with a great deal of uncertainty reflecting the ± continuous and often local variation in the relative lengths of carpophore and capsule.
There is, however, a distinctive morphotype characteristic of the populations from the coastal and nearinland parts of KwaZulu-natal and Swaziland.Plants from this region consistently produce stiffly erect stems, with closely branched, flat-topped cymes of flowers with short calyces, mostly 12-15 mm long, and unusually small capsules, 10-15 mm long.Although all of these states are found elsewhere in the species, their consistent association among these populations is unique and suggests some degree of genetic differentiation.We recognize these plants as subsp.polyantha.Distribution and ecology: throughout most of the range of the species but replaced along the eastern seaboard by subsp.polyantha (Figure 8).

Key to subspecies
Diagnosis and relationships: a variable taxon, typically with a lax, rounded inflorescence of relatively large flowers with calyx mostly 20-35 mm long, often with subulate lobes.The nature of the stem and calyx vesiture is very variable, with some plants entirely eglandular and others predominantly glandular, sometimes even within a single collection (e.g.Taylor 5180).Variation in the nature and degree of vesiture is especially evident among the populations along the southern Cape coast.Populations between Knysna and Plettenberg Bay often, but not always, have entirely eglandular stems and calyces.
Most populations have the anthers exserted beyond the mouth of the floral tube at anthesis, followed by the styles, which at anthesis are always included in the floral tube.This feature is thus evident only in older flowers.Also diagnostic of many populations are the long, subulate calyx lobes, 5-7 mm long but this can vary within populations.Plants with short calyces, ± 15 mm long, and carpophores ± 4 mm long, are typical of the West Coast, whereas specimens from western Lesotho (e.g.Schmitz 8095, Hilliard & Burtt 12065) have among the longest calyces (35-45 mm long) and carpophores (± 15 mm long) recorded for the species.
Plants are very variable in habit, those from coastal and alpine habitats typically more compact and prostrate whereas individuals from sheltered or shaded situations are more lax.Some of this variation is certainly ecological and linked to the habitat but some may well be genetic, since plants from the cliffs at Hermanus evidently retain their compact habit in cultivation [Davis s.n.(BOL)].
Separation from subsp.polyantha is not always clear in the Kwazulu-natal Midlands, especially around Pietermaritzburg, where plants have erect stems and wellbranched, flat-topped inflorescences but calyces ± 20 mm long.

Additional specimens
Distribution and ecology: largely restricted to the entrance to Saldanha Bay, where it is known from the breakwater and from across the lagoon in Postberg Reserve (Figure 10), but also known from shortly inland on the Farm Waterboerskraal east of Hopefield.The species is restricted to calcareous sands and consolidated calcrete dunes.
Diagnosis: a compact or cushion-forming perennial with erect or decumbent stems, distinguished by the mauve flowers with broad, overlapping petal limbs.The stamens and styles are included or shortly exserted but never protrude beyond the corona.Silene saldanhensis is probably most closely allied to the carmine-flowered S. ornata (Figure 11), which it resembles in stem vestiture and in the presence of scattered translucent testal cells in the seeds, but is distinguished by its broader, mauve flowers, colliculate seeds (Figures 2C, D), and in its ecology.Both species have stems with a mix of glan-dular hairs and short, scattered eglandular hairs (Figures 9B, 11B) but S. ornata is restricted to rocky habitats, and typically has a straggling growth form, even in very exposed situations, with the stems sprawling among bushes or over the rock surface.It typically has slightly smaller flowers, with petal claws ± 15 mm long and the limbs not overlapping, and tuberculate seeds (Figures 2E, F  The mix of glandular and short, eglandular hairs on the stems, and broad, overlapping petals, with included stamens and styles separate S. saldanhensis from coastal forms of S. undulata with a short calyx and carpophore.
Conservation note: the restricted occurrence of the species, the low number of plants and development at Saldanha render the species threatened.The Postberg population is protected as part of the West Coast national Park.

Additional specimens
WESTERn CAPE.-3317 (Saldanha): West Coast national Park, Kraalbaai, (-BB), 13 Sept. 2011, Manning & Goldblatt 3339 (nBg).3318 (Cape Town): Hopefield, Waterboerskraal Farm, (-AB), 3 Oct. 1974, Hugo s.n.(nBg).Distribution and ecology: a local endemic of the limestone hills flanking the mouth of Saldanha Bay (Figure 10), occurring on rocky slopes among coastal scrub.Plants mostly occur on the slopes or summit of hills, with the annual stems straggling through surrounding shrubs, but a population at Stony Head thrives on exposed limestone pavement directly facing the ocean.The plants here are very compact and almost cushionforming, with the flowering stems sprawling along the rock sheets.

Silene ornata
Diagnosis and relationships: Silene ornata is readily recognized by the unscented, deep carmine flowers that remain open thoughout the day and night.The branching of the flowering stems is largely asymmetrical, and stems and branches are densely puberulous, with a mix of short, glandular hairs and non-glandular hairs.The stamens are always included and the styles mostly so but in some plants they are exserted up to 4 mm beyond the petal claws.This is the form that is illustrated in Curtis's Botanical Magazine 11: t. 382 (1797).Other species in sect.Elisanthe have nocturnal, white to pale pink or mauve flowers that are fragrant at night.Silene ornata is evidently most closely allied to S. saldanhensis, another local endemic of Saldanha Bay, which shares a similar stem vestiture but which is restricted to calcareous sands and consolidated dunes, has slightly larger, mauve flowers with broader tepals, a more tufted habit, and colliculate seeds.
Distribution and ecology: restricted to the coastal forelands of the southwestern Cape, from Saldanha Bay along the West Coast to the Cape Flats, and at Vermont near Hermanus (Figure 10), occurring in deep, calcareous sands in strandveld thicket.
Diagnosis and relationships: although overlooked as a distinct species until now, Silene rigens is readily distinguished in sect.Elisanthe by its characteristic growth form and stem vestiture, flowers, fruits, and seeds.The stems are tall, stiffly erect, and largely unbranched except in the inflorescence, where the branching is symmetrical and the branches relatively short, resulting in distinctive, condensed triads of flowers and fruits.Flowering stems are strictly annual, dying off at the end of the season, to be replaced by axillary shoots from the base.The stem vestiture comprises only gland-tipped hairs, without an admixture of eglandular, acute hairs.Individual flowers are relatively small for the section, with white, dull pink or light mauve petals, the claws 8-10 mm long, and the anthers and styles ultimately barely exserted.The capsules, which are unusually broadly urn-shaped at maturity, 15-18 × 9-10 mm, and with a very short carpophore ± 2.5 mm long, cannot be fully accommodated within the calyx, which splits longitudinally at least midway but often to the base.The distinctly echinate seeds (Figures 2g, H Aug. 1978, Low 624 (nBg).3418 (Simonstown): Cape Flats, Swartklip, along road to military hut in E portion, (-BA), 22 Sept. 1972, Taylor 8211 (nBg, PRE).3419 (Caledon): Vermont, near the sea, deep sand, (-AC), 4 Sept. 1986 : 573 (1995b).[Chowdhuri's (1957)  geophytic perennial or suffrutex, mostly 10-50 cm tall; tuber parsnip-like or ovoid; stems decumbent or erect, ± suffrutescent below, basal woody portion 1-2 mm diam.but 0.5-1.5 mm diam.at base of inflorescence, ± densely puberulous-scabridulous with short, deflexed or spreading, acute hairs, rarely sub-glabrous.Distribution and ecology: widely distributed through the more temperate parts of southern Africa but absent from the arid interior, through eastern Africa northwards to the highlands of Sudan and Ethiopia, and extending across the Red Sea to Arabia (Figure 13).Silene burchellii favours loamy or fine-grained clay soils, often in rocky or stony places, from near sea level up to over 4 000 m.In southern Africa it is mostly a grassland forb but in the winter rainfall region occurs in open, mostly drier, fynbos and succulent shrubland.
Diagnosis and relationships: Silene burchellii is distinguished from the other southern African members of sect.Fruticulosae by its mostly erect or suberect stems, 0.5-1.5 mm diam.at the base of the inflorescence, and mostly linear to oblanceolate, rarely obovate leaves, (10-)15-60 × 1-16 mm.The leaves are ± concentrated in the lower part of the stem and well separated from the inflorescence by a distinct, leafless upper part ±100-150 mm long bearing one or two pairs of bract-like leaves.The calyx is very variable in length, 10-25 mm long, and is often slightly upcurved.The carpophore ranges from 4-12 mm long, varying from slightly longer than to shorter than the urn-shaped capsule, 8-10 × 5-7 mm.
Silene burchellii is sometimes confused with S. aethiopica in the Western Cape, but is distinguished from this annual species by its perennial, geophytic habit, and also never develops the highly branched growth form evident in well-grown plants of S. aethiopica.Although the tuber is seldom present on herbarium specimens, the relatively thick stem, terminating abruptly where it has been broken from the tuber, is characteristic.The two species are also ecologically separated, with S. aethiopica mainly coastal in deep sandy soils and S. burchellii essentially an inland species on finer-grained clay soils.Along the eastern seaboard, especially the Wild Coast and KwaZulu-natal, broad-leaved S. burchellii subsp.multiflora can be confused with S. crassifolia subsp.primuliflora.The latter can usually be identified by its prostrate or decumbent, often branching stems, mostly 1.5-2.0mm diam.at the base of the inflorescence, leathery or thick-textured leaves, somewhat expanded and strongly pleated calyx, mostly larger coronal scales, 0.8-1.0mm long, and broadly urn-shaped to subglobose capsules, (8-)10-12 × 7-10 mm.Its strictly coastal distribution, on sandy foredunes, is also diagnostic.
As currently circumscribed, Silene burchellii is the most widely distributed of the African species, ranging through the length of the continent, from Arabia and the uplands of Sudan and Ethiopia southwards though East Africa to the Cape Peninsula (Wickens 1976).Although highly variable in leaf shape and width, development of indumentum, number of flowers, and in calyx length, the species has resisted all attempts at segregation into more finely circumscribed taxa.The variation is often broadly correlated with geography and/or ecology but morphological patterns are confounded by the existence of numerous intermediates.The situation was summarised several decades ago by Turrill (1954), who concluded that it was reasonable to assume that the variation in S. burchellii is associated both with genetic and with environmental differences.This opinion has not altered since then, and Hedberg (1954Hedberg ( , 1957) ) was unable to maintain any afro-alpine infraspecific taxa on the morphological evidence without supporting evidence from further breeding experiments.
Despite the intractability of the species to satisfactory subdivision, workers on the tropical Africa material have consistently commented on the evident differences between the narrow-leaved tropical plants and the type of the species from the southwestern Cape, which is characterized by ± prostrate or decumbent stems and broad, oblanceolate to obovate leaves (Turrill 1954(Turrill , 1956a;;Wild 1961;gilbert 2000).There is a general feeling that the tropical African representatives should be distinguished at some level from all or at least some of the southern African material, and Wild (1961) accordingly treated all collections from the Flora zambesiaca area as var.angustifolia Sond.(1860) to separate it from typical var.burchellii from South Africa.With this signal exception, Sonder's (1860) early attempt at segregating the South African material of the species into four varieties, based largely on leaf shape and growth form, has not been implemented.The recent treatment of the species in Ethiopia and Eritrea by gilbert ( 2000) avoided all recognition of formal varieties in favour of five informal 'forms'.

A B
The southern African collections display extensive variation in vegetative and floral morphology but, as with the tropical African material, this is ± continuous, with no clear separation between the extremes, and we are unable to recognize more than a single species.There are, however, some general correllations between morphology and geography that we interpret as demonstrating some level of genetic differentiation within the species.Subsuming all of this variation under a single name obscures its existence and we therefore propose the recognition of four subspecies to highlight the most distinctive variants.The occurrence of intermediates between the subspecies makes the assignment of some specimens difficult, but a combination of morphology and geography is adequate for most material.Although not perfect, this classification provides a basis from which to work.
The type and other collections of Silene burchellii from the Cape Peninsula and adjacent coastal regions in the extreme southwestern parts of the Western Cape have a distinctive facies that sets them apart, notably the ± prostrate or decumbent stems and often broad leaves, and we treat these plants as comprising a narrowly circumscribed subsp.burchellii that is geographically isolated from other Western Cape populations by the coastal Cape Fold Mountains.A second set of populations from the coastal and near inland parts of the northern Eastern Cape and KwaZulu-natal and extending into Swaziland is distinctive in its broad leaves and multiflowered inflorescences of nodding or half-pendent flowers with very short calyces, mostly 10-12 mm long.We treat these plants as subsp.multiflora.A third set of populations with relatively long calyces, typically 20-25 mm long, is characteristic of the western, southern and central parts of the subcontinent, occurring along the western and southern Escarpment and the mountains of the southwestern Western Cape, inland onto the interior plateau.These populations are treated as subp.pilosellifolia.The remaining southern African plants have moderate-sized flowers, the calyx ( 12) 15-18(-20) mm long, and are treated as subsp.modesta.As accepted here, this subspecies is widely distributed, from the Mpumalanga Escarpment southwards into the KwaZulu-natal Midlands and through the foothills of Eastern Cape and KwaZulu-natal Drakensberg, and northwards into tropical and north Africa.
Although Carl Thunberg seems to have been the first to collect the species, during his visit to the Western Cape in 1772-1775, he mistakenly associated his collection (UPS-THUNB 10780) with the European species S. nutans L. (Table 1).Inexplicably, however, this name was omitted from his Flora capensis (Thunberg 1794), and the species was only recognized as distinct several decades later (Otth 1824) Bocquet & Kiefer: 8 (1978); SAM [2 sheets]!, S!, isolecto.).
Distribution: coastal regions of the Western Cape between Darling and Agulhas, and possibly at Port Elizabeth in the Eastern Cape, occurring mainly on shale or loamy soils in renosterveld but also in fynbos, on sandstone or limestone (Figure 14).On the Cape Peninsula, subsp.burchellii is found only on Signal Hill and Table Mountain, and is replaced by subsp.pilosellifolia on the southern peninsula.
We include Paterson 2543 (PRE) from Port Elizabeth here although it represents the only collection of subsp.burchellii from the Eastern Cape.The plants have prostrate, well-branched stems, oblanceolate leaves, and moderately sized flowers with calyx ± 15 mm long.These features are anomalous for subsp.pilosellifolia, the local form from the Port Elizabeth region.
Diagnosis: distinguished by the decumbent, divaricately branching stems with densely pubescent, obovate to oblanceolate leaves rarely more than 4× as long as wide; and moderately long calyx, 15-20 mm long.Plants from gansbaai and Agulhas are more compact and densely leafy than usual, possibly in response to their proximity to the ocean.
Silene thunbergiana, based on plants collected essentially in the same place as S. burchellii, was thought to differ in its broader leaves but this apparent distinction has no merit.Zeyher 240 (nBg, neo.!, designated here; nBg!, SAM [5 sheets]!, iso.).[The holotype appears to have been destroyed (Bocquet & Kiefer 1978) and we accordingly designate a neotype from the same geographical area as the type and that matches both the protologue and the current application of the name.Another collection that has been identified as type material by the Stockholm herbarium, Mund & Maire s.n.S-G-8699 (S), was collected at Voormansbosch near Swellendam and thus has no direct link to the protologue at all.] Silene burchellii var.angustifolia Sond.: 128 (1860).Type: South Africa, [Western Cape], Caledon Zwartberg, Dec. [without year], Ecklon & Zeyher 1959 (S, lecto.!, designated here; PRE!, SAM!, isolecto.).
Distribution and ecology: widely distributed through eastern southern Africa, Lesotho and Swaziland northwards into tropical and north Africa and Arabia (Figure 14), essentially in temperate or subtropical grassland and savanna, often in rock outcrops.
Diagnosis: subsp.modesta constitutes the core of this variable species after the more extreme forms have been segregated and includes ± suffrutescent, tufted plants with mostly oblanceolate to linear leaves and suberect, moderately-sized flowers with calyx (12-) 15-18(-20) mm long.This taxon seems to have included the bulk of the collections that were to have been recognized as a new species under the unpublished manuscript name Silene australis D.Masson ms.
Subsp.modesta is replaced in southern Mozambique and coastal KwaZulu-natal and Eastern Cape by subsp.multiflorus, mostly with broader leaves and with generally shorter flowers, the calyx mostly 12-15 mm long, and in the western, southern and eastern interior of the subregion by subsp.pilosellifolia with mostly longer flowers, the calyx 18-25 mm long.Subsp.burchellii, with procumbent or decumbent stems, is restricted to the extreme southwestern coastal regions.
The tropical African material, all with a calyx length of 12-20 mm, is comfortably included in subsp.modesta.Previously aberrant populations from northern Kenya, eastern uganda, and Ethiopia with exceptionally long calyces, 26-35 mm long, treated as var.gillettii Turrill (1954Turrill ( , 1956a)), have since been segregated as the distinct species Silene gillettii (Turrill) M.g.gilbert, also distinguished by its broader leaves with broadly cuneate or subcordate bases (gilbert 2000).Distribution: coastal and near inland from the Fish River Mouth in Eastern Cape northwards through Kwa-Zulu-natal into southern Mozambique and inland to near Mbabane in Swaziland, occurring mainly in subtropical grasslands on loamy or sandy soils (Figure 14).

Additional southern African specimens
Diagnosis: distinguished by the shortly decumbent or ± tufted habit with relatively broad, oblanceolate to obovate lower leaves, and the dense, multi-flowered inflorescence (typically with a well-developed branch), of small, nodding or slightly pendent flowers with short calyces, 10-12(-15) mm long.
The distinction between subsp.multiflora and subsp.modesta is blurred in the KwaZulu-natal midlands, where the transition between subsp.multiflora with broad-leaves and subsp.modesta with narrowly oblanceolate leaves, mostly less than 5 mm wide, lies along the Drakensberg foothills at an altitude of ± 1 500 m.
Coastal forms of subsp.multiflora with more decumbent branches may be difficult to separate from Silene crassifolia subsp.primuliflora in southern KwaZulunatal and Eastern Cape where the distribution of the two taxa overlaps.Introgression between the two species is one possible cause.An alternative hypothesis is that S. crassifolia is a coastal dune derivative of S. burchellii, with the two forms becoming progressively more distinct to the south as ecological and geographical isolation between them increases.Plants with the following combination of characters are treated as subsp.multiflora: stems not strongly prostrate, branching angles acute; inflorescence branched and multi-flowered with more than five flowers, well separated from foliage by leafless stem ±100-150 mm long bearing one or two pairs of bract-leaves; calyx 10-12(-15 mm) long; capsules ovoid, 8-10 × 5-7 mm.
South African material of this taxon has traditionally been identified as var.latifolia Sond.but this name is typified by a specimen from gauteng and correctly applies to broad-leaved forms of subsp.pilosellifolia.It was to have been recognized at species level by Daniel Masson, and several herbarium specimens bear the unpublished name Silene natalensis D.Masson ms.
Diagnosis and relationships: Silene crassifolia is usually readily identified by its prostrate or decumbent, relatively robust, perennial and often branching stems, mostly 1.5-2.0mm diam.at the base of the inflorescence, that are leafy almost to the base of the inflorescence, oblanceolate to obovate or sub-orbicular leaves, somewhat inflated and strongly pleated calyx 3-5 mm diam., and broadly urn-shaped to subglobose capsules, (8-)10-12 × 7-10 mm.The strictly coastal distribution, on sandy foredunes, is also diagnostic.
Collections of subsp.primuliflora from the eastern coast, especially the Wild Coast and KwaZulu-natal, may be confused with broad-leaved Silene burchellii subsp.multiflora and have sometimes been identified as a coastal form of this species.S. burchellii, which generally occurs on finer-grained soils inland of the coast, is a more slender plant, with essentially annual, mostly ± erect stems 0.5-1.5 mm diam.at the base of the inflorescence, with the foliage leaves concentrated towards the base of the stem and well separated from the inflorescence by an upper portion of leafless stem, and smaller capsules, 8-10 × 5-7 mm.This is the only region where both species occur, and the possibility of introgression in the coastal grasslands here cannot be discounted.
The two subspecies of Silene crassifolia recognized here have traditionally been regarded as distinct species but the greatly increased collections now available have led us to reconsider this interpretation.When Sonder (1860) last monographed the genus, he knew S. crassifolia only from the Cape Flats (Blouberg and Riet Valley [gordon's Bay]), and S. primuliflora from a few collections east of Knysna (Tsitsikamma, Port Elizabeth, and southern KwaZulu-natal).The distinction between the two taxa was quite clear from this sparse material: S. crassifolia was readily diagnosed by its suborbicular or obovate, hairy leaves, ovate bracts, hirsute calyx and relatively shorter carpophore half as long as the capsule, and S. primuliflora by its narrower, spathulate, scabrid leaves, ovate-lanceolate bracts, pubescent calyx, and relatively longer carpophore ± as long as the calyx.Later workers (Bocquet & Kiefer 1878;Masson 1989) have maintained these distinctions but the additional collections, especially those from the intervening southern Cape coast, show that these differences are not consistently maintained and that the two extremes grade into one another, especially between Knysna and Mossel Bay, where plants with the broad leaves and short carpophore of S. crassifolia develop the short pubescence characteristic of S. primuliflora.Further west, at Betty's Bay, plants have the narrow leaves of S. primuliflora but the woolly stem and calyx of S. crassifolia, while at Blouberg Strand some collections have the broad leaves and bracts of S. crassifolia but the short pubescence of S. primuliflora.Other purported differences between the taxa, in calyx length and capsule shape, are similarly variable across the range, with no clear discontinuity between them.This general but incomplete association of morphology with geography is consistent with the rank of subspecies (Stuessy 1990).

Additional specimens
Ecology and distribution: originally described from Plettenberg Bay, where it is still known only from the type and thus possibly extinct locally, the species is now also known to occur near Bredasdorp, more than 200 km to the west, in the De Hoop nature Reserve immediately west of Potberg (Figure 10).Here it is restricted to limestone outcrops near the coast, either fringing seasonal vleis or directly facing the sea.Plants growing on exposed cliffs display the typical maritime adaptations of a more compact habit, forming ± prostrate closely leafy mats, and with smaller, thicker leaves than those sheltered in potholes, which are more diffuse, forming loose cushions in the crevices.The species appears to be locally common where it occurs at De Hoop.
Diagnosis and relationships: one of the least known of the South African species, represented in herbaria by just a handful of specimens.
The highly reduced inflorescences of Silene mundiana, not immediately identifiable as monochasia, led Sonder (1860) to place the species provisionally ['doubtfully'] in sect.Elisanthe but its geophytic habit, eglandular vestiture, ovate calyx lobes, and especially the winged seeds place it correctly in sect.Fruticulosae.Here it is distinguished by its densely and closely branched habit, small leaves 7-15 × 1-3 mm, and ± filiform branches with just 1 or 2 flowers per inflorescence.Plants form small mats or low tangled mounds, with the numerous inflorescences extending only shortly above the vegetative growth.
Conservation note: the restricted occurrence of the species and its possible extinction around Plettenberg Bay render it vulnerable.
Distribution and ecology: widely distributed through the southern African winter rainfall region, mainly on the coastal lowlands, from the southern Richtersveld southwards to Stilbaai and inland to the Tankwa Karoo and Laingsburg, with a disjunction to Port Elizabeth in the east but evidently absent from the Little Karoo (Figure 18).The species is mostly encountered in welldrained, coarse-grained soils of a wide variety of types, ranging from acidic sands and gravels derived from sandstone and granite to basic limestones and calcareous sands, very rarely in fine-grained clays, occurring on sandy flats, in dry river courses, around salt pans and on limestone, or rarely dolerite, ridges.
Diagnosis and relationships: the only annual species of Silene native to southern Africa, S. aethiopica is distinguished from the introduced annual species, S. gallica and S. nocturna, by its wholly eglandular pubescencethe stems with short, deflexed hairs and the clavate calyx covered with short, appressed hairs-and by its discoid, peripherally winged seeds.Above ground it is superficially similar to the perennial S. burchellii (sect.Fruticulosae), with similar vestiture, calyx, and seeds and the two species can be confused in the southwestern parts of the Western Cape where their ranges overlap.Wellgrown specimens of S. aethiopica with branched stems are unmistakable but unbranched specimens without roots may pose problems.Calyx length is a useful discriminating feature in the region of overlap, with the calyx of S. aethiopica typically 10-15 mm long, except in subsp.longiflora from the Cold Bokkeveld, where it is 15-18 mm long.In contrast, the more common forms of S. burchellii in the region, subsp.pilosellifolia, have a calyx 18-25 mm long, although clayx length is shorter, 15-20 mm long, in subsp.burchellii in the extreme southwestern coastal areas.Ecologically, S. aethiopica prefers well-drained, sandy soils, whereas S. burchellii is mostly found in stony places or in finer-grained clays.The two taxa are, however, best separated by habit.S. aethiopica is an annual species with a slender taproot that is invariably present in herbarium specimens, which are always pulled from the ground with the main root intact.S. burchellii, in contrast, is a perennial with annual flowering stems but a persistent, woody stem base, developing a persistent, short or long rootstock that is swollen and tuber-like distally, although the swollen terminal portion is very rarely present in herbarium specimens.
As in other species of Silene, S. aethiopica is variable in stature, leaf morphology and flower size, notably in the length of petal limbs, carpophore and calyx.Much of the variation is either ecological or recurrent through the range of the species but populations from the southern Knersvlakte and the Olifants River valley are consistently less pubescent than usual, with the leaf blades either glabrous, apart from the margins, or glabrescent, and the stems often glabrescent as well.Large-and small-flowered forms occur throughout the range of the species, sometimes within single populations.
Strand forms of Silene aethiopica from the southwestern coast (Figure 17D), distinguished from more inland plants (Figure 17A, B) by the typical maritime adaptations of a more spreading habit, broader and more fleshy leaves, and fleshy calyx with distinctly bulging nerves, were described as a separate species under the name S. dewinteri (Bocquet 1977).The distinction between the two forms is not absolute, however, and some collections comprise material that could be assigned to both taxa (eg.Goldblatt & Porter 13509).We visited the type locality of S. dewinteri at Table View and confirm Bocquet's observations regarding the texture of the leaves and calyx but not of the habitall of the plants that we encountered were erect, possibly because our visit to the site was earlier in the season than Bocquet's.More significantly, however, our transect across the dunes passed through a gradation from typical S. aethiopica on the landward side to S. dewinteri on the seaward side.We encountered similar  Although plants at the two extremes of the ecological spectrum are certainly very different in appearance, they grade into one another in the ecotone, leading us to conclude that S. dewinteri represents a ± distinct ecotype rather than a separate species.This interpretation is also implicit in Bocquet's (1978) subsequent description of S. dewinteri as a 'sand-dune ecotype'.
That there is some genetic basis for the maritime form seems to be borne out by Bocquet's (1977) observation that the greater fleshiness of the plants was retained in cultivated plants grown from seed.It is not clear, however, whether this variant represents recurrent selection for a maritime genotype, a situation that is readily envisaged in a self-fertile species.In any event, we do not recognize this maritime form at any taxonomic rank.An investigation of the population genetics of the species is necessary before the taxonomic and evolutionary status of these variants can be accurately assessed.
A more clear-cut morphological disjunction is evident in populations from the Cold Bokkeveld and adjacent Cedarberg-Bidouw Mtns, which have markedly longer calyces (and carpophores) than usual.We describe this variant as subsp.longiflora.1791), which becomes a later synonym.Burman's description of the leaves of this species as viscid must be assumed to be an error as the type material does not show this feature.Curiously, the same specimen had earlier been determined as S. dewinteri Bocquet (1977) by Daniel Bocquet in 1976, who should therefore have taken up the epithet aethiopica for this taxon were it to be recognized at species level, which we do not.In any event, we regard the Burman specimen to represent a slightly luxuriant individual (possibly cultivated) of typical S. aethiopica and not the maritime 'dewinteri' form.
Although the source of Burman's specimens is not given, other collections of South African material in the Burman herbarium are known to have been made by [Heinrich] Oldenland, who was was employed by the Dutch East India Company at the Company gardens in Cape Town in the late seventeenth century.Ray (1704) certainly used material collected by Oldenland in the preparation of his Historia plantarum, and his mention of the species is a clear indication that it had reached Europe by the early eighteenth century.Linn. 583.8 [LInn, lecto.-digital image!, designated by ghafoor: 91 (1978)].
The species was first collected in South Africa under the name Silene ?pendulaL. by Carl Thunberg (Table 1), who travelled in the country from 1772 to 1774.Another early South African record of Silene nocturna that we have seen was made by Carl Zeyher (Zeyher 1960) in the mid-nineteenth century, somewhere in the vast tract of country between Malabarshoogde [near Queenstown in Eastern Cape] and Hessaquaskloof [near Riviersonderend in Western Cape].The next collection, from Wynberg Hill, dates from the end of the nineteenth century.Certainly, within two decades of this latter collection, the species was being found at several other localities around the Peninsula but has so far failed to establish itself as invasive.These early collections were misidentified as either S. gallica or S. clandestina [now S. aethiopica] or were left unnamed.
Diagnosis and relationships: the occurrence of Silene nocturna in southern Africa has been overlooked until now due to confusion with S. gallica, another introduced annual.The two species are superficially similar vegetatively and can be confused as herbarium specimens, although S. nocturna usually has more slender and lax flowering stems, typically with the lowermost flowers on long pedicels up to 35 mm long in fruit.They are best distinguished by details of stem vestiture, and especially by their flowers and seeds.Both species have a mixture of short and long eglandular hairs on the lower portions of the stem but in S. gallica the longer hairs are distinctly shaggy and 1-2 mm long, whereas in S. nocturna they are shorter and at most up to 1 mm long.The flowers of S. nocturna, as the name suggests, are typically nocturnal with distinctly bilobed petals (Figure 19C), and the calyx is appressed-puberulous, without glandtipped hairs and with only slightly longer hairs on the veins (Figure 19B).S. gallica, in contrast, has diurnal flowers with distinctive, unlobed petals (Figure 21C), and a glandular-haired calyx with characteristic glassy bristles 2-3 mm long on the veins (Figure 21B).Annual to 40 cm tall; stems ascending or erect, simple or well-branched from base and above, 0.75-3.00mm diam., lower parts closely puberulous with very short, ± deflexed or spreading hairs mixed with longer, shaggy, spreading or upcurved hairs 1-2 mm long but later glabrescent; upper parts of stems and inflorescences closely puberulous with very short, ± deflexed or spreading hairs mixed with short, spreading, gland-tipped hairs, inflorescence axis especially densely gladularpubescent.Leaves oblanceolate to spathulate-oblanceolate becoming oblong-lanceolate distally, 15-50 × 4-15 mm, apiculate, base narrowed in lower leaves, sparsely or densely pubescent with erect or ± appressed hairs, mixed on upper leaves with gland-tipped hairs, margins towards base ciliate with long, straggling hairs to 3 mm long, without evident side veins.Inflorescence a secund, spike-like monochasium terminating main stems and branches, (4-)8-15(-20)-flowered, lower flowers often remote; bracts smaller than leaves, unequal, fertile bract linear, sterile bract larger and more leaf-like, densely puberulous with gland-tipped hairs mixed with acute, eglandular hairs; pedicels mostly up to 3 mm long in lower (rarely up to 12 mm long in lowermost flower developed in axil of upper most leaf) but upper flowers subsessile.Calyx sub-cylindrical in flower, 8-10 × 1.5-2.0mm at anthesis, equally 10-veined without reticulate venation, densely puberulous with mix of gland-tipped and acute hairs and with conspicuous, glassy bristles 2-3 mm long on veins, lobes awl-like, 2-3 mm long.Flowers suberect at anthesis, unscented, diurnal.Petals longitudinally twisted and propellar-like, white or pink, sometimes with large crimson blotch or stain, claw 6-8 mm long, oblong, without auricles, limb obovate or oblong, minutely crenulate, unlobed, 1.5-4.0× 1.0-2.5 mm, coronal scales ± 1 mm long.Stamen filaments unequal, shorter series ± 4-6 mm long, included, longer series ± 7-9 mm long, exserted up to 2 mm, puberulous.Ovary pyriform, ± 2 mm long; styles ± 1.5 mm long, deeply included.Capsule ovoid, 6-8 × 4-5 mm, ± 6-8 × longer than carpophore, minutely transversely rugulose; carpophore ± 1 mm long, pubescent.Seeds 0.7-1.0mm, reniform, greyish brown, face concave-excavate, testal cells radially elongated-fusiform and mostly 2-4 granulate, back flat or weakly concave, tuberculate.Distribution and ecology: native through Europe and western Asia; now introduced as a weed into many parts of the world, and well established in South Africa, mainly in the winter-rainfall region, where it is most common in the southwestern Cape, ranging northwards onto the Bokkeveld Escarpment and the Kamiesberg in central namaqualand, and eastwards along the Eastern Cape coast to grahamstown and East London, with isolated records further north, in southern and central ; it is also recorded from the the highlands of east tropical Africa in Kenya, Tanzania, and uganda (Turrill 1956a).S. gallica typically occurs as a weed of agricultural lands and in waste places, along roadsides, and in other disturbed sites, but in some places has also invaded more open native vegetation that has been subjected to light disturbance, such as trampling and grazing.
The species was already established in South Africa by the late eighteenth century, when it was encountered here by Carl Thunberg, who travelled in the country from 1772 to 1774.Another early collection made near Port Elizabeth by Ecklon in the 1830s was misidentified as S. cernua Thunb., a later synonym of S. aethiopica Burm.
Diagnosis and relationships: an erect or somewhat spreading, rarely ±prostrate annual with obovate-spathulate lower leaves, densely glandular-puberulous on the upper parts of the stem and on the inflorescence, and highly characteristic flowers with ± entire petals.The calyx is ± tubular at anthesis with distinctive glassy bristles 2-3 mm long on the veins, and the urn-shaped capsule, 6-8 mm long, is carried on a very short carpophore, ± 1 mm long.The styles are very short, ± 1.5 mm long.
The seed testal cells are radially elongated on the flanks, which are deeply excavated, and each cell is ornamented with a single series of 2-4 granules (Figure 1E).A double series of cells along the midline demarcates the flat or weakly convex seed back (Figure 1F).
Minor variants have been distinguished in the past, based on branching, flower colour, petal shape, and differences in capsule orientation (greuter 1995a) but only the striking colour form, var.quinquevulnera, is recognized here.This variant cannot be distinguished from the typical form on any other grounds and identification of herbarium specimens depends on colour notes on the collecting labels or on residual coloration of the petals.Both varieties are sometimes represented on a single herbarium sheet.A second variety, var.anglica, was listed by Sonder (1860) as applying to more highly branched and spreading plants but as far as we can tell this is purely a manifestation of growing conditions and has no taxonomic value at all.11a.var.gallica Petals uniformly white to pink. Figure 21C.
Petals with large crimson spot or staining in centre of petal limb.Figure 21D.
Distribution: this variant occurs mainly in the southwestern Cape, extending into the Eastern Cape as far as grahamstown (Figure 20), but has not been recorded further east or north in southern Africa, nor from tropical Africa (Turrill 1956a;Wild 1961).It occurs mixed with the typical variety, usually in the minority.
Additional specimens *Collections comprising or including individuals of var.quinquevulnera.

FIguRE 5 .
FIguRE 5.-Recorded distribution of Silene vulgaris in southern Africa.
). Silene saldanhensis favours deep, loose sands or compacted dunes, forming neat cushions with the stems either decumbent or erect.It has mostly larger flowers with the claws 15-20 mm long and broad, often overlapping petals, and colliculate seeds, which are distinctive
Figure 13B.Distribution and ecology: widely distributed through western, southern and the eastern interior of southern Africa, from central namibia southwards along the western escarpment, through the Cape Fold Mountains into Eastern Cape as far as Zuurberg and inland onto the eastern interior plateau through Free State and gauteng as far north as Limpopo (Figure 14), occurring in a wide range of open shrubby or grassy habitats, mostly among rocks.
Jacobsbaai and at Elands Bay.

TABLE 1 -
South African Silene species in the Thunberg Herbarium (uPS-THunB).native South African taxa in bold.