Taxonomy of the genus Passerina ( Thymelaeaceae )

Passerina L. is mainly a southern African genus, comprising 20 species and four subspecies. A few species occur along the Great Escarpment, two extend into Zimbabwe and Mozambique, but most are concentrated in the Cape Floristic Region. Palynological. macromorphological and anatomical evidence was used in the delimitation of the genus and its infrageneric taxa. A cladistic study supports Passerina as a monophyletic genus. A genus treatment, key to species and a full species treatment are given. Each species treatment includes a taxonomic diagnosis, description and notes on taxonomy, etymolo­ gy, economic value and distribution. Illustrations of representative species are provided and distribution maps are included for each species. P. esterhuyseniae Bredenk. & A.E.van Wyk is newly described. A list of excluded species names high­ lights the previous cosmopolitan taxonomic interpretation of Passerina. as many names are now in synony my under other genera of the Thymelaeaceae.

The Thymelaeaceae is currently considered a family of ± 58 genera and ± 720 species.(Mabberley 1989;Brummitt 1992;Takhtajan 1997).It is subcosmopolitan and the distribution of the genera is listed by Mabberley (1989), as follows: Perhaps the economically most important character in the family is its tough fibrous bark.The bark of Wikstroemia, Daphne, Edgeworthia, Thymelaea and Daphnopsis is used for rope, and in the manufacturing of bank notes and strong paper.Flexible shoots of Dirca are used for baskets.Bark of Pimelea was used as a source of twine by early settlers in Australia.
Many genera are also known for their medicinal value.The wood of Wikstroemia is a source of incense and that of W. ovata C.A.Mey. is a strong purge.In China the bark of Daphne is used as an apparently safe and effi cient abortifacient: it contains the glycoside daphnin and an acrid resin (mezerein) giving plants a bitter taste.The decaying heartwood of Aquilaria malaccensis Lam. is saturated with a resin which is the basis of incense and when distilled it is used in perfume and medicine.
The genera Pimelea, Edgeworthia and Daphne are cul tivated for horticultural purposes.The scent of Daphne flowers is carnation-like and attractive to Lepidoptera; some members are moth-pollinated.Gonystylus bancanus (Miq.)Kurz. is a peat swamp-forest tree, with knee-roots.Its lightweight commercial timber is used for dowel ling and is much exported from Indomalesia.
In southern Africa, the bark of various genera is used for tying down thatch, for plaiting into whip thongs and for twine.Dais cotinifolia L. is an ornamental tree with attractive flowers, occurring mostly along the eastern regions of the country.

Passerina
In his comprehensive work on the circumscription of the Thymelaeaceae and infrafamilial taxa.Domke (1934) gave a complete historical review of the intergeneric clas sification of Passerina.He included the southern African genera Dais, Gnidia (= Lasiosiphon), Struthiola, Lachnaea (= Cryptadenia) and Passerina in the tribe Gnidieae, sub tribe Gnidiinae of the subfamily Thymelaeoideae.Breden kamp & Van Wyk (1996) place Passerina in the mono generic tribe Passerineae on the basis of mainly pollen characters.Currently Passerina is considered advanced at the intergeneric level, as many of the advanced character states present in other genera of the Thymelaeoideae are all found together in this genus.The most prominent charac ters distinguishing Passerina are the exserted.extrorse anthers and the unique anemophilous habit (Bredenkamp & Van Wyk 1996. 2001b).
The infrageneric classification of Passerina is docu mented by Linnaeus (1753) in his Species plantarum.in which he described P. filiformis, P. Iiirsuta, P. ciliata and P. uniflora.P. filiformis is the only species that is currently recognized in Passerina.Publications men tioned in the applicable protologue and in synonymy to the various species that pre-date the nomenclatural start ing point for the Spermatophyta [International Code of Botanical Nomenclature.Article 13.1 (Greuter etal. 2(KK))| are Linnaeus's Hortus Cliffortianus (1737), Van Royen (1740).Plukenet (1700: 180), Breyne (1678) and Burman (1739).The generic name Passerina appearing in Species plantarum (Linnaeus 1753) is associated with the subse quent description given in Genera plantarum (Linnaeus 1754) (Greuter et al. 2(XX).Article 13.4).Wikstrom (1818) recognized 41 species of Passerina and the subspecies P. filiformis subsp.divaricata: of these only tour species are presently recognized in Passerina.In the interim the subspecies was raised to species level and is presently known as P. falcifolia.Thunberg ( 1825a) recog nized nine species of which only one is currently main tained.His concept of P. glomerata, P. ericoides and Lachnaea conglomerata were completely incorrect and caused confusion right up to the present study.Meisner (1840;1857: 563-565) redefined the genus by clarifying 92 'species exclusae which were mostly synonymous with other cosmopolitan genera in the Thymelaeaceae and he retained only four species and six subspecies.The distribu tion of the remaining species clearly indicated that Passerina was a smaller genus, largely confined to southern Africa.At the beginning of the 20th century.Wright (1915) revised the Thymelaeaceae for the Flora capensis and his generic concept of Passerina was mostly based on that of Meisner (1857).He recognized ten species, of w hich three were new.as well as three subspecies.He recognized P. ericoides and Chymococca empetroides.We agree with Thoday (1924a) that C. empetroides is a synonym of P. ericoides.Although Thoday (1924a) provided a much improved classification of the group, the circumscription and identification of several species remained problematic, especially in the herbarium.Table 1 is a summary of taxa recognized in the most comprehensive works on Passerina from Linnaeus (1753) to the present study.
Etymology.Passerina refers to the Latin passer (= a sparrow) as the seeds resemble a sparrow 's beak.
Common names: the vernacular name 'sparrow-wort' was suggested by Miller (1768) for all Passerina species and Wendland (1798) used the name fadenformige Vogelkopf.According to Smith (1966) gonna is a collec tive name once used by the Khoekhoe for various mem bers of Thymelaeaceae, e.g.several species of Passerina and Struthiola.
Uses: many Passerina species grow on sand dunes and in sandy areas, with parts of the woody stem subter raneous, forming runners and developing an extended root system.Most of these plants are pioneers and resprouters, increasing their chances of survival in dis turbed areas.These plants are excellent sand binders and are suitable for binding problematic sandy areas, espe cially after the clearing of invader species.Sim (1919) recommended Passerina in his list of trees and shrubs for coastal areas exposed to sea winds.Certain Passerina species such as P. falcifolia are small trees and can be used as ornamental garden plants.P. filiformis has been cultivated in Britain and Europe since the time of Linnaeus.P. obtusifolia is used in the wild flower indus try in the Robertson area.The bark is exceedingly tough and is used for tying down thatch.According to Watt & Breyer-Brandwijk (1962) it is also plaited into whip thongs and used as twine.Members of the genus are not browsed by stock as the plants are apparently unpalatable (Story 1952).Ash from Passerina obtusifolia was tradi tionally used by the people of Genadendal in Western Cape in the home industry of soap-making.Although certain species have been recorded in cancer research, these plants are not currently known for their medicinal value.
Flowering and fruiting: most Passerina species flower profusely in spring, from September to October.During this season the Cape Floristic Region is quite windy and large amounts of pollen are produced, as Passerina is wind-pollinated.Pollen is often wafted away in clouds, causing a kind of hay-fever in sensitive persons (Marloth 1925).Fruiting time is mostly from December to January.The fleshy fruits of P. ericoides and P. rigida are dispersed by birds or rodents inhabiting the distribution ranges of these species along the South African coast.Fruits of P. truncata subsp.truncata, growing in the Karoo, passively fall to the ground, where they are probably dispersed by ants or rodents.The fruits of P. montana, occurring along the Great Escarpment.
are probably dispersed by birds as they are arranged at the tips of branchlets.exposed, red.and beak-like, possi bly resembling the beaks of nestlings.

Distribution and ecology:
in Passerina the highest number of species per grid (nine) occurs in each of the grids 3321 (Ladismith).3322 (Oudtshoorn) and 3419 (Caledon).The highest diversity of species (six) occurs in the False Bay area, from Seekoeivlei.including the Cape Flats, to De Mond at the Palmiet River (3418B).After an extensive study of herbarium material in co operation with field work.Western Cape is regarded as the centre of diversity for Passerina.from where certain species extend west, north and east.Thoday (1925) published an account of the geograph ical distribution and ecology of Passerina.based on 15 species.Of the 20 species currently studied, 10 are endemic and 4 species are near-endemic to the Cape Floristic Region.P. obtusifolia is widespread in the Northern.Eastern and W'estem Cape, whereas P. corym bosa occurs in Western and Eastern Cape, with outliers in KwaZulu-Natal.P. rigida is distributed from Western Cape, along the coast to northern KwaZulu-Natal; all these species are endemic to the southern African provinces in which they occur.P. drakensbergensis is endemic to the Bergville District in KwaZulu-Natal.P. montivaga is found from Mossel Bay and Oudtshoorn to Eastern Cape and along the escarpment northwards to Zimbabwe and P. montana is distributed from the east ern mountains and Great Escarpment of southern Africa to Zimbabwe and Mozambique.P. montivaga and P. montana are near-endemic to the Great Escarpment.
Nomenclatural notes: in the latter half of the eighteenth century Lachnaea conglomerata L. (1753), Passerina ericoides L. (1753) and P. glomerata Thunb. (1794) were constantly confused by botanists, causing Wikstrom (1818: 322) to place P. glomerata and L. conglomerata in the synonymy of P. conglomerata Thunb.In the same publication Wikstrom delimited and described P. paleacea.However, P. paleacea is not mentioned in Thunberg's revision of 1825, in which he described P. glomerata occurring in 'Hautbay', the currently known locality of both P. paleacea and P. ericoides.This confusion is re flected on many herbarium specimens, e.g. the specimen Herb.Thunberg 9579, bearing the inscriptions P. eri coides; P. glomerata (struck out) and the word 'paleacea' written in pencil.Although Thoday (1924b) chose the specimen Herb Thunberg 9597 as the type of P. paleacea, this specimen was not chosen as lectotype in the present study, as the Sparrman specimen cited by Wikstrom (1818: 324)  will always be doubt whether it is a duplicate of the Sparrman specimen cited by Wikstrom.

Diagnostic characters and relationships:
Passerina paleacea may easily be confused with P. rigida, as both occur on sand dunes along the coast.The branches of P. rigida are nodding and abundantly covered by pendulous branchlets, spikes are extended and the fruits are fleshy, yellow berries.Plants of P. paleacea are less robust, reaching a maximum height of 1.5 m. and are character ized by an abundance of subcapitulate inflorescences and dry fruit.The subcapitulate inflorescences at times led to the confusion of P. paleacea with P. truncata (= P. glomerata), but, these two species are morphologically as well as geographically distinct.P. paleacea has a mari time habit and P. truncata is distributed from Vanrhynsdorp, along the Cederberg Mountains, to Malmesbury, Ceres, Tulbagh and Matjiesfontein up to Seven Weeks Poort.The earlier confusion between P. paleacea and P. ericoides was probably due to their sympatric occur rence, but these two species are morphologically quite different.
Distribution and ecology : Passerina paleacea occurs in both the Southwestern and the Agulhas Plain Centres of the Cape Floristic Region (CFR) (Goldblatt & Man ning 2000) and is a typical fynbos element.It grows on coastal dunes and in maritime habitats from Langebaan, round the Cape Peninsula to the Cape Flats.Kogel Bay, Hermanus, Gansbaai.De Hoop, the Potberg coast, Bredasdorp, Arniston.Vermaaklikheid and Puntjie up to Stilbaai (Figure 2).The vegetation types dune fyn bos and dune thicket form a mosaic along many parts of the southern Cape coast (Lubke 1998a(Lubke , 1998b;;Lubke & Van Wijk 1998).This same distribution pat tern is displayed by P. paleacea as it is found in the dune scrub, amongst typical fynbos species, but not in the dune thicket amongst larger shrubs or small trees with mesophytic leaves such as species of Chrysanthemoides, Mimusops, M orelia, Rhus and Sideroxylon.
Nomenclatural notes: according to the concept of Wright (1915), P. ericoides is not only distributed along the southern coast of Western Cape (present interpreta tion), but also along the coast to Eastern Cape and further inland up to Mpumalanga.However, most of the inland specimens cited by him have been classified as P. mon tana by Thoday (1924a).The interpretation of P. corym bosa by Wright (1915) posed the same problem, as Wood 4036 (the lectotype of P. montana) was also placed in this taxon.Meisner (1857) described P. rigida var.tetragona cit ing two Drege specimens, one from Ezelsbank and the other from Stormberg.The Ezelsbank specimen (Drege 2971, P, K) is P. truncata, but the Stormberg specimen could not be located.According to Gunn & Codd (1981), Drege crossed the Stormberg (3126BC, Queenstown) on 17 December 1832.The first author suspected that the Drege specimen would be P. montana, as it is common in this area.This suspicion is supported by Sim 68 (from the Pirie Mountains in the King William's Town District), a syntype of P. montana, bearing the inscription 'P.rigida Wiks-tetragona' and the Drege specimen from Storm berg is consequently regarded as P. montana.Hilliard & Burtt (1988) noted two rather distinct forms of P. mon tana in KwaZulu-Natal.The first form is characterized by plants on rock platforms that are low, rounded bush es, 0.3-1 m high, with the tips of the branches erect, whereas those of the second form inhabit valleys and are riverside bushes of up to 2 m high, with open branches and pendulous branchlets.The present study, taking the whole distribution range of P. montana into considera tion, recognizes two forms.One, centred in the God's Window area of Mpumalanga, are rounded shrubs 0.5-2 m high, with many branchlets covered with smaller, de cussate, imbricate leaves, bluish green in colour.The second form dominates in the Free State.Lesotho, KwaZulu-Natal and Eastern Cape.These plants are more robust, with open branches and larger, yellowish green leaves and inflorescences, which are tinged pink.However, the two forms are not geographically distinct and intermediates are common.Both forms unequivocal ly show the specific characters and therefore we do not propose to give them formal taxonomic recognition.Studies V of the leaf epidermis, anatomy and floral morphology (Bredenkamp & Van Wyk 2000, 2001a, 2001b) supplied no further evidence on which the two forms could be delineated.

Diagnostic characters and relationships:
Etymology: the specific epithet is derived from the Latin montanus (= pertaining to or growing on moun tains).This is a very appropriate epithet as P. montana is distributed along the Great Escarpment from Eastern Cape to Zimbabwe.
Common names: Cooper 2302 (K), from Lesotho, re ported the vernacular name Likhabei and Staples 17 (PRE), from the Maluti Mountains in Lesotho, recorded the name Lekaphu.Story (1952)  Distribution and ecology: Passerina montana is a near-endemic to the Great Escarpment of southern Africa, with distant satellite populations in high mountain areas of Angola, Namibia and Limpopo [Northern Province], South Africa.It is distributed from Nyanga in Zimbabwe, along the escarpment to Manica and Sofala in Mozam bique, Limpopo, Mpumalanga, Swaziland, KwaZulu-Natal, Free State, Lesotho and the Eastern Cape (Figure 5).Outliers in Angola have been found on the escarp ment of the Huilla Plateau near Lubango and the Cheila Mountains.Several specimens of this species have been collected at Moltkeblick on the Auas Mountains in Nami bia.In Limpopo, P. montana is found in the Soutpansberg area and on the Blouberg, as well as on the summit of Krantzberg in the Waterberg Mountains.A single speci men (Goossens 375) was collected in the Pretoria District, but the species is currently probably extinct in this area, due to human impact.This species grows at altitudes of (900-)l 200-3 000 m.At Nyanga, P. montana is associated with Erica mannii and E. hexandra, bordering on Brachystegia woodland and montane forest.In Mozambique and South Africa it has been found with Widdringtonia nodiflora and Erica species, bordering on montane forest.It is common amongst rocks on hills, mountain slopes, mountain tops, cliff ledges and rocky ridges.It also frequents stream courses and banks as well as riverbeds and banks, where the growth form has been reported as a shrub amongst rocks, a drooping bush over running water, a limply spreading bush in sand or dense bushes.These plants also grow in river valley forests and along plantations.
Diagnostic characters and relationships: bearded sepals, leaves and bracts distinguish this species from P. pendula.
Eponymy: this plant was named in honour of the explorer and botanist W.J. Burchell, who collected in Caledon and as far north as Tulbagh between 1810 and 1811.During this trip Burchell 7761, the lectotype of P. burchellii, was collected on the summit of the mountains of Baviaanskloof near Genadendal.

Distribution and ecology:
Passerina burchellii is en demic to the Southwestern and Langeberg Centres with in the CFR.It is common on mountain summits of the Villiersdorp and Genadendal Districts (Figure 7), with outliers on southeastern rocky slopes of Towerkop in the Swartberg Mountains at Ladismith.This species occurs at altitudes of 1 333-2 167 m, often covered in mist.It is found in small groups on sandy loam, between boulders and rocks on upper south-or southeast-facing slopes.Nomenclatural notes: in the Catalogue o f the Linnaean Herbarium.Savage (1945) made the following inscription 'Tulb.list c. 1769.n.l.det.L.-Blaeria eri coides'.This refers to consignments of bulbs, seeds and herbarium specimens that Rijk Tulbagh sent to Van Royen, the Burmans at Amsterdam and Linnaeus at Uppsala (Gunn & Codd 1981).Jackson (1917Jackson ( , 1918) ) published a list of 203 of the specimens sent to Linnaeus around 1769 and identified by him.The first inscription on the list is the provisional name Blaeria ericoides, which Savage (1945) believed to be the P. ericoides specimen at LINN, but there is no numbering or any other indication on the specimen to link it with Tulbagh's list (Jackson 1917(Jackson -1918)).As Linnaeus had already described P. ericoides in 1767.the specimen at LINN is probably not part of the Tulbagh collection.Thoday (1924a) clearly regarded the specimen at LINN, named by Linnaeus, as the type of P. ericoides.As no other origi nal elements exist, P. ericoides LINN 504.5 is regarded as a lectotype designated by Thoday (1924a).
Thunberg ( 1825a) accepted Wikstrom's concept of P. glomerata, occurring at Hout Bay in the Cape, and cited P. ericoides in synonymy, causing confusion about the identity of the latter taxon.Meisner (1840) reinstated P. ericoides, but the concept of this taxon became even more doubtful in the light of the cited distribution.In 1857 Meisner retained his concept of P. ericoides.occur ring at Uitenhage.Port Elizabeth, Witbergen and Onderbokkeveld.and placed P. glomerata and Lachnaea conglomerata in synonymy under P. ericoides.This revi sion by Meisner (1857) was largely followed by Wright (1915).Because of his incorrect concept of the taxon.Meisner (1857) was confronted with material from Table Bay and Standvallei with red berries, which he then named Chymococca empetroides, based especially on the fleshy fruit.Thoday (1924a) was justified in placing this name in synonymy under P. ericoides, as the descriptions of these taxa coincide and as the fleshy fruit of C. empetroides is not unique, but is also found in P. rigida.The concept of P. ericoides.occurring along coastal dunes mainly in the Cape Peninsula and adjacent coastal areas of the Western Cape, was clarified by Thoday ( 1924a) and is also accepted in the present study.
Diagnostic characters: Passerina ericoides is charac terized by greenish flowers, with a coriaceous, strigose hypanthium and the fruits are fleshy red berries.The leaves are greyish green and oblong, with an obtuse apex.The bracts are leaf-like, larger and lanceolate.
Etymology: the specific epithet ericoides refers to the ericoid appearance of this species indicated by the phrase 'corollae tubus globosus, inflatus-unde et Ericam refert flore', which was used by Linnaeus (1767) in his origi nal description of the species.
Common names: Willdenow (1799) introduced the vernacular name heideartiger Vogelkopf and the com mon names 'Christmas berry* or dronkbessie were docu mented by Smith (1966).
Uses: Marloth (1925) remarked that P. ericoides was laden with bright, scarlet fruits and that it was often employed as a Christmas decoration.The juicy pulp has a somew hat unpleasant taste, but appears to be harmless (dronkbessie).As early as 1919.Sim recommended P. ericoides as a useful shrub for planting in coastal areas exposed to sea winds.This species occurs on coastal dunes and on the banks of lagoons in the Cape Peninsula and adjacent coastal areas of Western Cape.The plants are excellent sand binders as they have an extensive root system from which resprouting often takes place.Because human impact and invasion of alien vegetation along the coast of the Cape Peninsula are very high, reha bilitation and conservation of coastal dunes is of vital importance.P. ericoides plants are ideally suited to com bat erosion of coastal dunes and can be used as a substi tute in coastal areas where alien vegetation is cleared.In their research on the coastal erosion of the Milnerton beaches.Many-stemmed, much-branched, robust shrubs of (0.6-)1.0-2.0(-3.04)m tall on coastal dunes; secondary and tertiary branches ascending, conical in appearance, formation of branchlets profuse, decussate, older branch lets self-pruning, lax or arcuate, 6G-100 mm long, pro gressively shortening towards growing point, young branchlets ascending, 5-60 mm long, growing point nod ding, fertile branchlets often pendulous and secund.Stems greyish brown, bark stringy; cork grey-brown; branchlets and growing points densely white-tomentose, tomentum forming lengthwise patterns with cork on older branchlets, which later become glabrous.Leaves imbricate, overlapping ± 50%, appressed, plane shape lanceolate to ovate, length x depth 1.6-2.5 x 0 .1-1.1 mm, adaxial surface concave, villous, abaxial surface convex, glabrous, greyish green, smooth, often covered by salt crystals; base sessile, dilated; median vein in dis tal half visible as a keel, forming acute apex; margins vil lous.Inflorescences with spikes usually extended, 6-10flowered, arrangement subterminal, axis white-tomen tose, proliferating growth common.Bracts appressed, ascending in fruit, widely ovate, length x depth (2.6-) 3.4 x 1.4(-1.9)mm; lamina adaxially concave (inside), abaxially convex (outside), villous inside, glabrous out side, smooth, with 2 or 3 shallow folds on each side of main vein, wings absent, greyish green, coriaceous; base dilated; apex with distinct, short, acute point; margins tomentose, involute.Floral envelope ± 4 mm long, mem branous and yellow during pollination, dehydrated after shedding of pollen, turning red.Hypanthium glabrous at ovary, neck tomentose, ± 0.8 mm long.Sepals: outer sepals cymbiform, midrib adaxially tomentose, abaxial surface glabrous, inner sepals obovate, adaxially tomen tose, abaxially glabrous.Androecium with filaments of antipetalous whorl ± 0.5 mm and those of antisepalous whorl ± 1.5 mm long; anthers, ± 0.8 x 0.4 mm, sub-basifixed, 2-thecous and 4-locular.Ovary 2.2 x 1.4 mm.Fruit a fleshy yellow berry, ± 2.6 x 2.3 mm, enveloped by persistent, loosely arranged hypanthium, fragmenting over widest circumference of fruit, the fragmented hypanthium, sepals and androecium being shed.Seed ± 1.4 x 1.1 mm. Figure 8.
Nomenclatural notes: in his description of P. rigida, Wikstrom (1818) clearly indicated the specimen of Sparrman, in the Thunberg Herbarium, as the type.This specimen bears the inscriptions Passerina glomerata p and epithet rigida in pencil.Thoday (1924a) identified the handwriting of the pencilled 'rigida' as Wikstrom's, comparing it to signed letters in the library at Kew. Wikstrom's handwriting was also confirmed in the pre sent study, using examples published by Burdet (1979).According to Stafleu & Cowan (1986), original speci mens of the Thunberg Herbarium (to which Sparrman also contributed) were donated to UPS and the duplicates D were sent to S. Thus three other Sparrman s.n.specimens of Herb.Swartzii, Herb.Wikstromii and Herb.Gastromii, housed at S, and a fourth one from Schreber's herbarium, housed at M, are duplicates.As Wikstrom clearly indi cated the specimen in the Thunberg Herbarium as the type, we regard it as the holotype and the other four Sparrman specimens as isotypes.
Diagnostic characters and relationships: Passerina rigi da is easily distinguished as a robust, rigid shrub, usually 1-2 m high.The ascending branches are conical in shape due to many branchlets that are pendulous when fertile.The flowers are yellow and membranous and bright yel low berries are borne subterminally.The leaves are nar rowly lanceolate to ovate and the apex is acute, with the main vein visible as a blunt keel.The bracts are widely ovate with the apex acute.This species is easily distin guished from P. paleacea which occurs on secondary dunes and is distributed mainly along the southern coast of Western Cape.Plants of the latter species are less robust, reaching a maximun height of 1.5 m, character ized by an abundance of subcapitulate inflorescences and the fruits are dry (achenes).
Etymology: the epithet rigida refers to the rigid, ascend ing branches, characteristic of the growth form of this plant.
Uses: Passerina rigida is a pioneer of the coastal dunes along large portions of the South African coast.Because these robust plants are excellent sand binders and are completely adapted to maritime winds and salt spray, they can be used in the rehabilitation of coastal dunes in disturbed areas.P. rigida has an extensive root system from which resprouting commonly takes place.
The yellow berries are an important food source for ani mals inhabiting coastal areas, especially birds.
Distribution and ecology: Passerina rigida is distrib uted from Witsand River Mouth on the western coast of the Cape Peninsula, along the coastline of South Africa to Lake Sibayi on the northeastern coast of KwaZulu-Natal (Figure 9).It is endemic to the coastlines of KwaZulu-Natal, Eastern Cape and Western Cape.The specimen Taylor 4143, recorded as far North as Lambert's Bay on the West Coast, is regarded as an out lier, as no other specimens have been recorded in the grid 3318.Thoday (1924a)  This species occurs on littoral sand dunes and ham mock dunes just above the level of spring tide.It is also found in marshy places and on sandy banks of river mouths and lagoons.A stunted form is present on shal low marine sand over limestone and on rocky hills fac ing the sea.Lubke & Van Wijk (1998) regard P. rigida on the southern and Eastern Cape coast as a pioneer found in bush clumps or bush pockets on rear dunes.According to them, there are often no pioneer communi ties on the vast dune sands and the first vegetation encountered as one moves away from the shore is dune thicket, in which P. rigida is one of the dominant shrubs.Passerina species occurring on littoral dunes in Western Cape are found mainly in Coastal Fynbos (Acocks 1988).From the southern Cape coast to Port Alfred, dune fynbos and dune thicket form a mosaic as well as a successional series between the two vegetation types (Lubke & Van Wijk 1998).For a complete description see Bredenkamp & Van Wyk in Bothalia 32: 76-79 (2002c).Distribution (Figure 9).

Conservation status: Lower Risk [LR-lc] (Victor 2002).
This species is rare, but does not qualify for Red List sta tus under IUCN (2000)   Shrubs or shrublets 0.3-0.5 m high.Stents greyish brown, younger branchlets greyish tomentose; cork fine ly fissured, grey-brown, displaying whitish sclerenchy-ma fibres at scars.Leaves imbricate on young branchlets, closely appressed to stem, cymbiform.plane shape lin ear-lanceolate, length x depth ± 2.0 x 0.5 mm; lamina inversely ericoid.adaxial surface concave, setose, abaxi al surface convex, glabrous; base sessile; apex rounded into subacute point; margins sometimes ciliate.Inflores cences polytelic synflorescences; main florescences and co-florescences spicate.Bracts enveloping flowers and fruits, largest after anthesis of flowers, becoming more coriaceous and rounded at fruit set.decussate, imbricate, sessile, helmet-shaped, widely obovate in outline, length x depth t 3.1 x 2.4 mm: lamina adaxially concave (inside), abaxially convex (outside), setose on inside, glabrous on outside, thinly chartaceous.smooth, concolorous.brownish, extending into a membranous rim or membranous wings; base cuneate; main vein extending to form a subacute to acute apex: margins ciliate in dis tal half.Floral envelope constituting hypanthium (fused calyx and androecium) and sepals; membranous and yel lowish during pollination, dehydrated after shedding of pollen, becoming papyraceous, turning red to brown, ± 4.6 mm long.Hypanthium a membranous cylindric tube, indumentum at ovary and neck tomentose, neck ± 0.7 mm long, abscission tissue and articulation plane absent.Sepals 4, petaloid.imbricate in bud.flexed in flower: outer sepals concave oblong with apex adaxially tomen tose, abaxially setose: inner sepals concave, obovate with apex adaxially glabrous, abaxially setose.Androecium: filaments of antipetalous whorl ± 0.4 mm and those of antisepalous whorl ± 1.2 mm long.Ovary ± 1.8 x 0.5 mm.Fruit enveloped by persistent, loosely arranged hypanthium fragmenting over widest circum ference of fruit, the fragmented hypanthium.sepals and androecium being shed; an achene with pericarp mem branous and dry.± 2.5 x 1.2 mm. Figure 10.Diagnostic characters and relationships: Passerina esterhuyseniae is easily distinguished from P. comosa by its helmet-shaped bracts, which are widely obovate in outline.The concolorous, brownish bracts are thinly chartaceous and smooth in texture, the lamina extends into a membranous rim or membranous wings and the main vein elongates forming a subacute to acute apex.The flowers are membranous and yellowish during polli nation and red to brown after shedding of the pollen.
Eponymy: this species is dedicated to Elsie Esterhuysen who diligently collected plants especially the high-mountain flora of the Northern, Western and East ern Cape.

Distribution and ecology: Passerina esterhuyseniae
has been collected on the northern Cederberg Mountains at Groenberg near Pakhuis and at Konpoort (Figure 11).It is endemic to the Northwestern Centre within the CFR.The northern Cederberg area is covered by Mountain Fynbos (Rebelo 1998).This species grows at the peaks of mountain tops at altitudes of ± 1 167 m, or against rocky slopes amongst high rugged rocks.Confined most ly to mountainous areas, this species is still undercollect ed.Pillans 7689 (BOL) collected on slopes near the road SE of Redelinghuis has been classified under P. ester huyseniae, although these plants seem to be more robust and grow at lower altitudes.
Diagnostic characters and relationships: Passerina comosa and P. quadrifaria both have abaxially hairy bracts and are easily confused.However, these two species are geographically segregated, with P. comosa considered as a 'north-western endemic' of the Cape flora (Weimarck 1941), whereas P. quadrifaria is endemic to the Karoo Mountain and Southeastern Centres.Mor phologically P. comosa is less robust, intemodes are longer, leaves adhere closely to the stem and are gener ally more hairy and the bracts often have extended wings that are abaxially setose to villous.
Etymology: the epithet comosa refers to the hairs on the abaxial surface of the leaves, bracts and sepals, which are characteristic of this species.
Distribution and ecology: Passerina comosa ranges from mountain summits and slopes of the Kamiesberg to Calvinia in the Northern Cape (Figure 11).In Western Cape it is distributed in the area between 33° and 34°S lat itude and from 19° to 21°E longitude, with Primos 41 (PRE) as the most easterly outlier.This species is endem ic to the Northern Cape, as well as the Northwestern, Southwestern and Karoo Mountain Centres within the CFR.It occurs on the Roggeveld, Witteberg and the Klein Swartberg Mountain Ranges.This species is found in sand among rocks, on rocky ledges, on mountain summits, or on SW-facing slopes at altitudes of 1 000-1 200 m.13.
Nomenclatural notes: Passerina pendula.ascribed to Ecklon & Zeyher.was first published as a nomen nudum by Drege (1847b).This name was placed in synonymy under P. rigida var.comosa by Meisner (1857).Wright (1915) partly followed Meisner s interpretation of P. rigi da var.comosa.but in the citation of the specimens he added all those that were later published as P. burc hellii by Thoday (1924a).In his revision of Passerina.Thoday ( 1924a) reinstated the name P. pendula Eckl.& Zeyh.ex Meisn., as the varietal name 'comosa* had already been used at species level by Wright (1915).The present study Diagnostic characters and relationships: Passerina pendula is distinguished from P. burchellii by being taller (up to 1.5 m), much-branched shrubs with pendu lous branchlets, with grey-green, softly coriaceous and smooth leaves and yellow-pink membranous flowers that are abaxially glabrous and adaxially scantily tomentose.
Etymology: the specific epithet pendula refers to the pendulous branchlets of these shrubs as seen in their nat ural habitat.

Distribution and ecology:
Passerina pendula is ende mic to the Southeastern Centre within the CFR.It is dis tributed on hills and slopes from the Kouga Mountains in Western Cape to the Langkloof Mountains and the Great Winterhoek Mountains in Eastern Cape (Figure 14).The species is also distributed along watercourses as it occurs in the KwaZunga Catchment Basin and on the banks of the Upper Swartkops River as well as the Bushmans River at Port Elizabeth.P. pendula grows at altitudes of (133-)383-600 m.On mountain slopes it is often found in a belt above valley thicket and below mountain fynbos.It grows in sand or shallow, gravelly, sandy loam.The plants are frequent throughout the nat ural range of the species and a number of populations are conserved in the Groendal Nature Reserve at Uitenhage.
Nomenclatural notes: Passerina galpini C.H. Wright (1915), published with a full description, but without a Latin diagnosis, was accepted by Thoday (1924a).In the present study the specific epithet is corrected to 'gal pin ii' and the name accepted as validly published by Wright, as the starting date for a Latin diagnosis as pre requisite for valid publication is 1 January 1935 (Greuter et al. 2000).

Diagnostic characters and relationships: Passerina
galpinii is distinguished by its characteristic bracts, which are oblate, with the midrib extended into a leaf like point; the lamina is cymbiform, greyish green, char taceous and glabrous, with the midrib adaxially tomen tose; the wings are straw-coloured, membranous, broad ly rounded and bullate.The distribution of this species is also diagnostic as it is endemic to the Agulhas Plain Centre within the CFR.
Eponymy: Passerina galpinii was named in honour of the botanist E.E.Galpin.The holotype of this name, Galpin 4491, was collected on 7 October 1897.At this time, Galpin organized a collecting trip from Port Eliza beth via the Humansdorp.Knysna.George, Riversdale, Swellendam and Caledon Districts to Cape Town and increased his collecting numbers from 3531 to 4846.All these specimens were probably identified at the Bolus Herbarium in Cape Town, where he also spent a few weeks (Gunn & Codd 1981).
Common name: Rebelo (1998) mentions the vernacu lar name Elim gonna for this species.

Distribution and ecology:
Passerina galpinii is ende mic to the Agulhas Plain Centre within the CFR.It is dis tributed on stony flats, coastal limestone deposits and limestone hills, from Elim to Bredasdorp.Amiston.Stilbaai, Melkhoutfontein, Albertinia and Mossel Bay (Figure 14); it grows at altitudes of 0-290 m.Plants reach a height of ± 1.2 m on stony flats, but become stunted on southeast-facing slopes of limestone hills, overlooking the sea.The plants are frequent in their natural environ ment.They are conserved in the De Hoop and Potberg Nature Reserves and several private nature reserves.The vegetation of the area is threatened by large stands of Acacia cyclops (rooikrans), an alien invasive tree.
Passerina galpinii is associated with Laterite Fynbos (Rebelo 1998), occurring on the Elim Flats of Western Cape, which is characterized by gravelly, lateritic and seasonally waterlogged soils.The present study also indicates the presence of this species in Limestone Fyn bos, where it occurs on coastal limestone deposits.

Diagnostic characters and relationships:
Passerina drakensbergensis is characterized by appressed leaves, up to 6.5 mm long.The bracts are lanceolate, up to 7 mm long, the apex is obtuse to acute without a leaf-like point and membranous wings are absent.It may easily be con fused with P. montivaga and P. montana, both occurring in the northern KwaZulu-Natal Drakensberg area.P. montivaga has longer leaves (up to 8 mm) and bracts with the midrib extended, forming a straight or filiform, leaf-like point.The wings are ovate with margins hairy in the distal half, or obtrullate, narrowing abruptly into the midrib.P. montana can be separated by its terminal subcapitulate spikes and short leaves (up to 4 mm long), which are linear to lanceolate, with a dilated base and with a prominent median vein in the upper third of the leaf, incurved at the acute apex.The bracts are ovate to obovate in outline.
Passerina montivaga is a fynbos element which pos sibly originated in the southern Cape and dispersed east wards.Both this species and P. montana are distributed from Eastern Cape via the Drakensberg Mountains northwards to Zimbabwe.Although the distribution of P. drakensbergensis, P. montivaga and P. montana overlap in the northern part of the KwaZulu-Natal Drakensberg, significant trends in the geographical and altitudinal ranges of these species have been identified.Etymology: the specific epithet refers to the location of this species in the northern KwaZulu-Natal Drakens berg.

Diagnostic characters and relationships:
Passerina corymbosa is distinguished by its greyish green leaves and grey-brown stems of which the older branchlets are leafless, often arcuate and indurate.The leaves are lateral ly compressed with the distinct midrib somewhat keeled.The rhombic to obtrullate (diamond-shaped) bracts are always conspicuously angled and distinctly 4-or 5ribbed.The most diagnostic character in the leaf anato my of P. corymbosa is the presence of a hypodermal sclerenchymatous sheath, illustrated by Bredenkamp & Van Wyk (2001a).This species has always been con fused with P. filiformis sensu lato.which has inconsis tently been distinguished by longer, filiform leaves.In Western Cape P. filiformis subsp.filiformis is separated by widely obovate bracts, narrowing abruptly into a fili form point.Where the distribution of P. corymbosa and P. montivaga overlap in the southern Cape, the latter species is distinguished by bracts with ovate wings and margins that are hairy in the distal half.In Eastern Cape P. montivaga is distinguished by obtrullate bracts nar rowing abruptly into a straight, leaf-like point.
Etymology, of all the species in the genus, Passerina corymbosa is the most common, as it is adapted to a wide range of habitats mostly in Western and Eastern Cape.The specific Latin epithet corymbosa (= with a cluster of flowers or of fruits) indicates the 10-16-flowered, extended spikes usually arranged in multiflowered main and co-florescences.Van Wyk & Gericke (2000).P. corymbosa (formerly known as P. vulgaris) is also called bakkersbos, a name that commemorates an era when the official bakers in the Cape used this plant to heat up their outdoor ovens.

Common names: according to
Uses: from an agricultural point of view.Story (1952) described P. corymbosa at Keiskammahoek as an un palatable bush, which remained undamaged from grazing, among the few closely cropped specimens of Cliffortia linearifolia and C. paucistaminea.However, the value of P. corymbosa as a pioneer, and also in combatting ero sion, cannot be underestimated.This species is common ly found along roadsides and in other disturbed places.It is one of the most successful species for the rehabilita tion of embankments along newly built roads in Western and Eastern Cape.The plants are resprouters from woody, underground rootstocks and are excellent sand binders, often found on coastal sand dunes.Considering the human impact and invasion of alien vegetation along the Cape coast.P. corymbosa would be a natural pioneer, combatting erosion in areas where alien vegetation is cleared.
Distribution and ecology: except for a few outliers, P. corvmbosa is endemic to Western and Eastern Cape, and all the phytogeographic centres within the CFR.Although this species is distributed from Clanwilliam to Cape Town and eastwards to East London, it most com monly occurs in an area between the coast and the 33°S latitude and from 18° to 29°E longitude (Figure 17).Gerstner 105 (PRE), collected near Compasberg in the Lady Grey District, represents the most northerly distri bution of P. corymbosa in Eastern Cape.The specimens collected in KwaZulu-Natal are regarded as outliers, rep resenting remnants of a former wider distribution.Hilliard 4081 (PRE), collected at the Ellesmere Farm in Ngome (KwaZulu-Natal), is an anomalous specimen, with a greyish appearance, infected by fungi and record ed from cliff faces.This specimen was classified as P. cory mbosa on the basis of the angular bracts and the leaves that are laterally compressed.The other two speci mens, Herb.Poeppig s.n.. probably collected before 1868, and Rudatis 1204 (PRE), collected in 1910, repre sent populations that have possibly succumbed to human impact.
Passerina corymbosa is a species with a wide habitat spectrum.It most commonly occurs as a pioneer along roadsides over the whole range of its distribution.The species is found in stony areas on mountain slopes, peaks and mountain passes.Along the coastal region, it is often found on the rear dunes.It also grows in river valleys and on the banks of river mouths.This species is common in the whole of the Fynbos Biome of the CFR.In Eastern Cape it is found in all the above-mentioned habitats, but also in grassland.Story (1952) reported that P. corym bosa is found in sourveld and mixed grassveld but that it showed no sign of advancing into the sweetveld.In open grassland this species is often clustered along streambanks or on rocky areas.P. corymbosa occurs at a range of altitudes, from sea level up to 1 300 m.
Diagnostic characters and relationships: the growth form of P. obtusifolia may easily be confused with that of P. corymbosa.but P. obtusifolia is distinguished by Etymology: the Latin specific epithet, obtusifolia, refers to the obtuse apices of leaves and bracts, which are char acteristic of this species.
Common name: the vernacular name karoo gonna is used by the local people at Genadendal.
Uses: according to the curator at the Museum in Genadendal, P. obtusifolia was traditionally used by the local people in the home industry of soap-making.The plants were burnt and the alkaline ashes used to react with the stearic acid in fat at boiling point, thus forming soap.In the Robertson area these plants are used in the wild flower industry.Bayliss 521 (PRE) is a voucher specimen recorded in cancer research, but the results must have been negative; these plants are not currently known for their medicinal value.
Distribution and ecology: Passerina obtusifolia is endemic to Northern.Western and Eastern Cape (Figure 19).It is centred in a belt between 33° and 34°S latitude and from 19° to 27°E longitude, comprising all the cen tres within the CFR-it is most common in the Karoo Mountain.Langeberg and Southeast Centres.ary of fynbos and karroid vegetation and is common in the Little Karoo, growing at altitudes of (300-)670-l 400 (-1 700) m.Although this species occurs at high altitudes on the summit of the Swartberg Pass, it grows below the snow line and does not occur on the highest peaks of mountain ranges in its distribution range.It is common in drier mountainous habitats, growing in shallow rocky soil and between rocks on well-drained slopes.It also grows amongst sandstone boulders of upper mountain slopes and on stony ridges of mountain tops.On the Hantamberg it has been recorded in renosterveld on the flat, rocky, dolerite summit.On Jonaskop it grows in a zone below the fynbos and is absent at the summit.This species is also found amongst rocks in river valleys and dry streambanks.The average height of these plants is 0 .8-1.8 m, but stunted forms have been recorded from the arid Bergkwagga National Park, which is one of the most northeasterly localities.P. obtusifolia is a very com mon species and amongst the dominant species within its distribution range.

Diagnostic characters and relationships:
Passerina paludosa is a stout shrub up to 2 m high, occurring most ly in marshy ground on lowland flats.It is characterized by erect, nearly straight, greyish green, imbricate, ap-pressed leaves, which are ± lanceolate.The bracts are narrowly obtrullate, with the midrib and leaf-like point stout and the apex acute.This species is distinguished from P. filiformis subsp.filiformis which has filiform leaves and widely obovate bracts, which narrow abrupt ly into a filiform point.
Etymology: the specific epithet paludosa refers to the habitat of this species, namely marshy lowland flats; from the Latin paludosus (= marshy, swampy or boggy).
Distribution and ecology: Passerina paludosa is endemic to the Southwestern Centre within the CFR (Figure 19).Herbarium specimens dated from 1921 to 1995 show that this species used to be distributed from sandy places along the Malmesbury Road (Acocks 24X2).along marshy areas of the Cape Flats and the Stellen bosch District to the Palmiet River at Elgin, the most easterly locality.As P. paludosa was severely affected by urbanization and invasion by alien vegetation in the Cape Peninsula, it is currently confined to small marshy areas east of Muizenberg.
According to Smuts (1996) the only three extant popu lations known, are at the Rondevlei Nature Reserve.Zeekoevlei and along the Strandfontein Road.Label information on Peterson 1263.collected in 1982, states that the population at a housing estate site SE of Zee koevlei consisted of ± 4(X) plants, but Smuts (1966) reported only 60 living plants.At the same time the popu lation at Rondevlei consisted of 35 plants and the one along the Strandfontein Road of possibly a few hundred.Currently both the Zeekoevlei and Strandfontein sites are in danger of urban development and are being threatened by invasive alien vegetation, primarily Port Jackson (Acacia saligna) and rooikrans (A.cyclops).Conserva tion measures proposed by Smuts (1996) include an environmental impact study at the Zeekoevlei site prior to any development and a plea for urgent attention by conservation authorities to ensure the conservation of the Strandfontein population.
The Rondevlei Nature Reserve boasts more than 250 plant species of which many are rare and endangered.Species associated with P. paludosa include Chondropetalum nudum.Juncus krausii and Leucadendron levisanus.In recent years the management at the reserve concentrated on restoring and managing its biodiversity.Alien vegetation has been cleared, plant species that occurred there historically have been re-introduced and P. paludosa has been successfully propagated by cut tings to expand the population.As aridification is an important effect of urbanization and as alien vegetation impacts on the natural drainage system of an area, the whole wetland east of Muizenberg can be conserved only if it is included in the Rondevlei Nature Reserve (Smuts 1996).
Recently two new populations of plants, that appear to be P. paludosa.were collected at Springfontein Farm near Stanford [3419AD.Louw 7083 (NBG.PRE)].and in seasonally wet clays at Heidehof.5 km NW of Pearly Beach [3419CB.Helme 2376 (NBG.PRE)].These speci mens were not included in the distribution of P. paludosa as further population studies need to be done.Taking urbanization and invasion by alien vegetation into account, the Red List status of P. paludosa was also not changed.
Nomenclatural notes: Wright (1915) overlooked the combination P. filiformis L. var.divaricata Wikstr. (1818), also indicated by Thoday (1924a), which is the earliest name for the taxon.However, this name based on Sparrman s.n.(Herb.Thunberg 9573) falls into syn onymy under P. falcifolia, as the name of a taxon does not have priority outside the rank in which it was pub lished (Greuter et al. 2000).

Diagnostic characters and relationships:
Passerina falcifolia is distinguished by the mottled grey-green fal cate leaves, which are inclined or horizontally spreading.The widely ovate bracts are villous inside and narrow into a leaf-like, falcate point, with chartaceous wings that are distinctly ± 4-ribbed and reticulately veined.The most conspicuous floral character is the slender, often arcuate, tomentose hypanthium neck, exserted from the clasping bract.This species may be confused with P. fili formis subsp.filiformiswhich has widely obovate bracts, narrowing abruptly into a filiform point.The bracts are basally to centrally setose on the inside and the wings are glabrous.P. montivaga is another close species, but is distinguished by bracts that are basally setose on the inside, with glabrous wings.
Etymology: the specific epithet falcifolia is derived from the Latin falcatus (= curved like a sickle), referring to the falcate or sickle-shaped leaves of these plants.Uses: Passerina falcifolia is used for fuel or for mak ing cord (Palmer & Pitman 1972).According to label information on Dahlstrand 1905 (PRE), the species is cultivated by florists.Plants grow into small ornamental trees and could be used more widely in horticulture.According to Grobbelaar 63 (PRE), P. falcifolia is a host to members of the insect genus Eremnus.Distribution and ecology: Passerina falcifolia is asso ciated with forests and Mountain Fynbos (Rebelo 1998) in the southern Cape and the southern parts of Eastern Cape.It is a near-endemic to the CFR and occurs in the Karoo Mountain, Southwestern and Southeastern Centres, as well as the Zuurberg, Blaauwkrantz and Alexandria Forests of Eastern Cape.It most commonly occurs in a belt between the coast and the 33°S latitude and from 22° to 26°E longitude (Figure 22).The two specimens.Much-branched, erect shrubs, with rigid branchlets and inflorescences or smaller, extensively branched, round ed shrublets under arid, calcareous habitat conditions, (0.2-)0.3-0.75(-l.1) m high.Stems: older ones grey-brown, indurate, and sclerenchyma fibres exposed; young stems reddish brown, indumentum whitish tomentose, forming lengthwise patterns with cork on older branches, which gradually become glabrous; cork Assuring lengthwise; intemodes longer than leaves during prolific lengthening of branchlets or shorter under arid conditions.Leaves greyish green, ascending, appressed.decussate and rigid, or under arid conditions, imbricate (overlapping 5-30%), appressed or ascending, diverging at an angle of up to 30°: lamina narrowly lanceolate or oblong, longitudinaly folded, triangular in section, length x depth 2.4-4.3 x 0.7 mm, adaxial surface concave, tomentose, abaxial surface glabrous; base sessile; apex obtuse; margins glabrous, involute.Inflorescences with conspicuous, multiflow ered main and co-florescences; spikes robust, rigid, extended, narrowly ellipsoid, with rows of enlarged, decussate, pointed bracts, 20-30-flowered.arrangement subterminal, axis white-tomentose, proliferating growth common.Bracts grey-green, rose-tinted during flower ing time, ascending, imbricate, widely ovate, midrib shortly extended into a point, length x depth (4.3-)5.1 x 1.8(-2.0 ) mm; older bracts folded lengthwise along midrib, younger bracts adaxially concave (inside), abax ially slightly convex (outside), villous inside, glabrous outside, coriaceous; wings widely ovate, chartaceous, ± 5-ribbed.reticulately veined; base cuneate; apex acute; margins ciliate in distal half.Floral envelope ± 8.4 mm long, papyraceous and yellow-pink during pollination, dehydrated after shedding of pollen, turning red to brown.Hypanthium glabrous at ovary, neck exserted.sparsely pubescent.± 2 mm long.Sepals: outer sepals cymbiform.ad-and abaxially glabrous, inner sepals obovate, adaxi ally scantilly tomentose, abaxially glabrous.Androecium with filaments of antipetalous whorl ± 0.7 mm and those of antisepalous whorl ± 1.7 mm long; anthers ovoid, ± 0.9 x 0.3 mm, sub-basifixed, 2-thecous and 4-locular.Ovary ± 2.7 x 1.1 mm.Fruit an achene with pericarp membra nous and dry, ± 2.1 x 1.2 mm, enveloped by persistent, loosely arranged hypanthium, breaking up at neck base due to dehydration and torsification of tissue, resulting in the sepals and androecium being shed.Figure 23.
Nomenclatural notes: as the starting date for a Latin diagnosis is 1 January 1935 (Greuter et al. 2000), P. rubra is a valid name, although it was published with a full description, but without a Latin diagnosis, by Wright (1915).The combination P. filiformis L. var.squarrosa (Meisner 1857), was overlooked by Wright (1915), but mentioned in synonymy by Thoday (1924a).In the pre sent revision all the type material cited by both Wright (1915) and Meisner (1857) was studied.Galpin 4492 (K) was selected as the lectotype of P. rubra by Thoday (1924b) and Zeyher 3779 in S was selected as lectotype for P. filiformis var.squarrosa as it is internationally available in many herbaria.
Diagnostic characters and relationships: the distribu tion of P. rubra partly coincides with that of P. corym bosa, P. montivaga and P. falcifolia.P. rubra is a small er shrub (average height 0.3-0.75m), often occurring in calcareous soil.It is distinguished from the other three species which are taller (average heights 1-2 m), and especially from P. falcifolia, which is a tall shrub or a small tree (up to 3.04 m), often associated with indige nous forests.P. rubra may also be separated by the inflores cences which have extended, robust spikes, with up to 30 fertile, enlarged bracts.The bracts are typified by the midrib which is shortly extended into a point and by the wings which are adaxially tomentose, widely ovate, char taceous, ± 5-ribbed and reticulately veined.Flowers are distinguished by the exserted hypanthium neck, which is ± 2 mm long and glabrous to sparsely pubescent.Etymology: the specific epithet rubra was derived from the Latin ruber (= red), referring to the conspicu ous, multiflowered inflorescences of these plants, which have 20-30 flowers arranged in four rows and turning red after wind pollination.
Uses: Passerina rubra is a pioneer which often occurs along roadsides or in disturbed places, e.g.close to the salt works in the vicinity of Port Elizabeth.It is also found on calcareous soils between Port Elizabeth and Cradock.In the Coega area, earmarked for industrial de velopment, P. rubra might be a useful plant for combat ting erosion.
Distribution and ecology: Passerina rubra is near endemic to the CFR, occurring in the Langeberg, Karoo Mountain and Southeastern Centres, as well as southern parts of the Eastern Cape.It most commonly occurs in a belt between the coast and the 33°S latitude and from 20° to 26°E longitude.P. rubra is distributed from the Bontebok National Park in the Swellendam District, eastwards to Gowie's Kloof near Grahamstown (Figure 22).This species is somewhat variable.It was initially thought that plants in Western Cape were more rigid, with longer intemodes and appressed leaves, which did not overlap, whereas those in Eastern Cape tended to be rounded shrublets, with imbricate, ascending leaves.After many specimens, from all parts of the range had been studied, no geographical or morphological discontinuity between the two forms could be shown, and it was decided that the morphological differences were probably due to plas ticity.Plants growing in more arid conditions and cal careous soil, typical of the Port Elizabeth and Cradock areas, tend to be rounded much-branched shrublets, with short intemodes and imbricate, ascending leaves.Under more favourable conditions in sandy loam, the plants are taller, less branched, intemodes are longer and the ap pressed leaves do not overlap.
Passerina rubra is common in the Steytlerville, Humansdorp.Port Elizabeth and Grahamstown areas of Eastern Cape and less frequent in Western Cape.The area be tween Cradock and Port Elizabeth is renowned for the ancient dunes and flats, abounding in limestone.Acocks (1988) described the vegetation occurring on the lime stone as False Fynbos (A70), also known as Mountain Fynbos or Grassy Fynbos (Rebelo 1998).P. rubra seems to be well adapted to the calcareous soils on which it occurs.These plants are often pioneers in disturbed areas and along roadsides, as in the Colchester.Coega and the Markman industrial areas of Port Elizabeth.At the Groendal Catchment Basin, this species occurs in grass land on sandstone and it is also found on semi-karroid.dry, rocky hillsides in the Baviaanskloof area.At the Bontebok National Park it is found in flat areas between fynbos species.P. rubra grows at altitudes of 70-7(X) m.
mentions the name pakaan.Von Breitenbach et al. (2001) used the names berg-gonna and mountain gonna.Uses: information on the specimen Watt & Breyer-Brandwijk 1851, collected at Thabaneng, states that the plants are used medicinally.However, Watt & Breyer-Brandwijk (1962) supplied no further details.
mentioned Bowker s.n.from Somerset, Cooper 2301 from Albany and Ecklon & Zeyher s.n.(SAM 19801) as specimens from inland local ities.In recent years more cases of P. rigida growing along sandy banks of rivers adjacent to the coast have been noted.

FIGURE 17 .
FIGURE 17.-Known distribution of Passerina drakensbergensis, O; P. corymbosa, ■ FIGURE 19.-Known distribution of Passerina obtusifolia, • ; P. paludosa, ■ Brown 25975 and Rogers 28858, collected near Caledon in October 1924, are regarded as outliers, possibly indi cating a wider previous distribution of the species into areas with woody vegetation in Western Cape.P. falcifo lia occurs from Meiringspoort, in the Oudtshoom area, to Ruytersbosch in the Mossel Bay area, and along the Outeniqua, Tsitsikamma and Great Winterhoek Mountains to the Grahamstown area.Passerina falcifolia is found on mountain plateaus and southeast-facing slopes on Table Mountain Sand stone in shallow, sandy loam soil.Plants commonly occur along forest margins, in open patches, or disturbed areas along roadsides.This species is also found in coastal regions and riverine fynbos.P. falcifolia grows at a range of altitudes, from sea level up to 1 100 m.Consenation status: Least Concern (LC) (IUCN Spe cies Survival Commission 2000).

TABLE 1 .
-A summary of taxa in the most comprehensive works on Passerina
truncata are found in the Karoo Desert National Botanical Garden at Worcester.P. obtusifolia usually occurs on the northern side of the southern Cape mountain ranges in drier habitats and P.falcifolia is con fined to the summits of mountains and southwards towards the southern Cape coast.Intermediates between P. obtusifolia and P. falcifolia have been found on the boundary between the two species, just north of the Prince Albert Pass.

-gonna, gonna has, gonna bos, kannabas and
Outeniqua gonna' or the Outeniekwagonna, refer ring to the Outeniqua Mountains where it occurs (Coates Palgrave 1977).Palmer & Pitman (1972) use the vernacu lar name forest gonna, as these plants are commonly seen along roadsides on mountain passes of the southern Cape forests.Von Breitenbach et al. (2001) use the names OuteniekwaOuteniqua gonna, gonna bush.