The genus Amphiglossa ( Gnaphalieae , Relhaniinae , Asteraceae ) in southern Africa

It is proposed that the generic concept of Amphiglossa DC. be broadened to include the genus Pterothrix DC. The two gen­ era were previously separated on the presence or absence of ray florets. A thorough study of various characters showed that the type of florets was overemphasised in the past and that the pappus and cypselas provide sufficient evidence to unite the two genera The genus consists of 11 species of which Amphiglossa celans Koekemoer. A. grisea Koekemoer, A. rudolphii Koekemoer and A. susannae Koekemoer are newly described A key to the species as well as detailed descriptions and distri­ bution maps are provided. Photographs are included for each species and key characters for the genus are also illustrated.


INTRODUCTION
Formerly the two genera, Am phiglossa DC. and Pterothrix DC., were poorly known and distinguished by the absence or presence of ray florets.Even De Candolle (1838) was not always sure to which genus certain species belonged, hence his description of A. perotrichoides DC. for a species without ray florets.A thorough study of various characters showed that the type of flo rets was overemphasised in the past and that the pappus and cypsela provide sufficient evidence to unite the two genera.The genus consists of 11 species, four of which are newly described here.
The extensive fieldwork I undertook over a period of six years during different seasons, to cover the different flowering times, was very valuable in revising this group.Not only were four new species discovered but another three, that were mainly known from type mater ial, were re-discovered after earlier collections by Ecklon & Zeyher and Drêge.
Nine species in the genus are characterised by very restricted distributions in Namibia and the Northern, Western and Eastern Cape of South Africa.Two other species, A. tom entosa (Thunb.)Harv.and A. triflora DC. have wider distributions in the other South African provinces and Namibia.Specimens were studied in BM.BOL.BR.E, G, G-DC.K, LINN, NBG.P, PRE, S. SAM.SBT, TCD, TUB.UPS and WAG (abbreviations as in Holmgren et al. 1981).Many of the older collections are of a very poor quality and were of limited value in the revision.
Perennial, woody, much-branched, wiry shrubs or dwarf shrubs, erect or decumbent, stoloniferous or rhizomatous.Branches glabrous or with a thin layer of arachnoid hairs, tips rarely spinescent.Leaves small, imbricate or widely spaced, often in fascicles, sessile or inconspicuously stalked, ericoid.convex, abaxial surface arachnoid or glabrous, adaxial surface woolly, margins smooth, tips mucronate.acute or rounded.Capitula radi ate or discoid, terminal, sessile, solitary, or in small groups of distinct units, homogamous or heterogamous; ray florets 3-12, bilabiate or pseudobilabiate florets female, disc florets bisexual.5-lobed.Involucral bracts in several series, outermost chaffy or foliaceous, inner most longer, narrower.Receptacle discoid, nude or with small extensions around cypsela bases.Anthers 5, syngenesious, apically acuminate, basally tailed.Style bifid, stigmatic area in two strips along margins of style arms, inflated cells at branch tips short, rounded.Nectary pre sent in all florets, a small disc between style base and cypsela.Cypselas small, asymmetrical, glabrous, papil lose.Pappus bristles 18-32, densely plumose from tip to base, in one series, free or rarely fused at the very base.Flowering time: mainly November-M arch, some species in spring-sum m er and others in summer-autumn.
Am phiglossa is recognised by its small, sessile capit ula that are terminally borne, solitary or in small groups, a chaffy involucre, papillose cypselas (Figure 1 A. B) and pappus of many fine bristles (Figure 1C.D) that are pro fusely plumose and interlocking in the central parts.Am phiglossa also has a germination strategy unique in the Relhaniinae.Fresh, mature cypselas germinate with in 24 hours by means of a basal hygroscopic substance that absorbs water very quickly, causing the seedcoat to burst and thereby facilitating germination.A gelatinous blob (Figure IE.F), about the size of the swollen cypsela, then surrounds the root tip.A m phiglossa callunoides DC., Prodromus 6: 259 (1838).Type: Eastern Cape.3325 (Port Eh/.abeth): District Uitenhage, in the stony channel of the Zwartkops River, fl.Jan. 1835, Ecklon 51 (G-DC.lecto.!, here des ignated).Figure 2B.
Stoloniferous, woody, multistemmed, blue-grey shrubs, to 0.5 m tall.Branches slender and very flexible, erect at first, bending over and becoming stoloniferous, rooting where they touch the ground; older plants form ing well-anchored bushes, covering an area of up to 6 m2; young active shoots up to 3 mm diam., older, main branches 1.5-2.0mm diam.Primary brachyblasts devel oping in every leaf axil on main stems, numerous sec ondary brachyblasts developing in leaf axils of primary brachyblasts; stems of brachyblasts and young branches often white-tomentose; brachyblasts mostly negatively geotropic, whereas main branches positively geotropic, thus creating the impression that secondary branching is restricted to one side of the main branch.Leaves persis tent, sessile, adpressed.narrowly triangular, margins entire, slightly inrolled, midrib prominent, adaxial surface woolly, abaxial surface smooth, tips shortly mucronate, young leaves yellowish green, older leaves fading to grey, giving a characteristic blue-grey sheen to plants; subtend ing leaves on main stems alternate, up to 10 mm long and 2 mm wide at base, about a leaf length apart, leaving branch exposed; brachyblast leaves minute, up to 2 mm long, imbricate, subtending leaves on brachyblasts not much longer than ordinary' leaves.Capitula radiate, heterogamous, 7-8 x 2-3 mm.8-10-flowered with the same number of ray and disc florets; single head terminally borne on brachyblast, linear when young, involucre spreading at maturity to release pappus and seeds.Outer involucral bracts 10-15.foliaceous in upper part and along midvein, remaining green around midrib, in 3-6 series, ovate.2.5-4.5 mm long, increasing in length from outer to inner series, the outer about half the length of innermost bracts.Inner involucral bracts scarious along margins, with multicellular glands.10-15, in 4 or 5 series, 6.0-7.5 mm long, linear to narrowly oblong, hya line towards apex, those in innermost series hyaline in upper half, mucronate.as long as pappus.Receptacle flat tened, less than 2 mm diam.. honeycombed, ridges short er than 1 mm.Bilabiate or pseudobilabiate florets white, female, 4 or 5; lamina ± 1.5 x 0.5 mm, 3-lobed; tube 4.5-5.0mm long.Disc florets white, bisexual, 5-lobed.5.0-5.5 mm long.Cypselas subterete, ± 1 mm long, chestnut-coloured, inconspicuously ribbed, papillose, developing in 3-5 florets, more often in bilabiate or pseudobilabiate florets; released in a cluster together with pappus at maturity.Pappus bristles 18-23, ± 5 mm long, barbed in lower fifth, plumose in upper part, seta up to 1.5 mm long.Flowering time: January-April.
Diagnostic characters: stoloniferous growth form, numerous brachyblasts; leaves very small; capitula small, terminal on brachyblasts; small white florets just visible at tip of involucre.
Distribution and habitat: known from two localities only: a short section of the Kwa-Zunga River (inland part of Swartkops Riser), NW of Uitenhage between Springfontein and Rooikrantz Farms, and the Farm Hoeree in the Kouga Mountains, NW of Joubertina (Figure 3).Observed growing in abundance amongst boulders in the bed of the Kwa-Zunga River.ish or brick-red with a shiny appearance, often with zigzag branching pattern.Leaves sessile, ascending, elliptic to linear, margins entire, inrolled, twisted, adaxial surface woolly, tips shortly mucronate.up to 8.0 x 1.5 mm, seemingly caducous after growing season, greyish green, very sparse on most branches, ± half a leaf length apart on terminal branches; young, active shoots up to 3 mm diam., large-leaved, leaves flat, up to 16 x 6 mm.Capitula discoid, homogamous, solitary in last 3 or 4 ter minal leaf axils, 7-8 x 2-3 mm, 5-flowered.Involucral bracts 12-17, up to 8.0 x 1.3 mm, stereome clearly visi ble in central upper part, outermost bracts foliaceous, joined together by hairs on outside, innermost bracts scarious along margins, tinged dark wine-red towards slightly recurved, mucronate apex.Receptacle conical, less than 1 mm diam., honeycombed, ridges shorter than 1 mm.Florets white or pink, bisexual, 5-lobed, 4.5-5.0mm long.Cypselas subterete.± 1 mm long, inconspicu ously ribbed, papillose, annulus not distinct.Pappus bristles 18-26, connate at very base, ± 4 mm long, barbed in lowermost part with barbs interlocking, short ly plumose along shaft ending in tufted tip, seta up to 0.2 mm long.Flowering time: February-April.
Distribution and habitat: poorly known, currently only from a small area between Garies and Kotzesrus (Figure 3).'Strandveld', deep sand, with sparse vegeta tion. of predominantly mesembryanthemous and asteraceous elements that seldom grow taller than 500 mm.Erect, rhizomatous, chaffy-green shrubs, up to 0.4 m tall.Branches rigid, up to 4 mm diam., secondary branches ascending, to 2 mm diam., developing in each leaf axil on main stem; older plants forming wellanchored bushes; stems becoming rhizomatous and pro ducing copious shoots when covered by sand deposits during floods.Brachyblasts developing in each leaf axil of secondary branches, generally short, up to 5 mm long, bearing capitula close to stems; secondary brachy blasts rare.Young branches and stems of brachyblasts often white-tomentose.Leaves bright green, sessile, adpressed.triangular, margins entire, slightly inrolled, midrib prominent, adaxial surface woolly; subtending leaves to secondary branches on main stems alternate, up to 11 mm long, 4 mm wide at base, ± a leaf length apart, leaving branch exposed, browning with age and then often reflexed; subtending leaves on secondary stems much shorter, up to 5 mm long, 1.5 mm wide at base, becoming brown in older parts, ascending to inclined; leaves on brachyblasts minute, up to 2 mm long, imbricate.Capitula radiate, heterogamous, 7-8 x 2-3 mm, 8-10-flowered.with same number of ray and disc florets; single heads terminally borne on short brachyblasts, linear when young, involucre spreading at maturity to release pappus and seeds.O uter involucral bracts 10-15.2.5-4.5 mm long, foliaceous in upper part and along midvein, in 3-6 series, ovate, increasing in length from outer to inner series, the outer about half the length of innermost bracts.Inner involucral bracts scar ious along margins, 10-15, 6.0-7.5 mm long, in 4 or 5 series, linear to narrowly oblong, those in outer series remaining green around midrib, hyaline towards apex, those in innerm ost series hyaline in upper half, mucronate, as long as pappus.Receptacle flattened, less than 2 mm diam.. honeycombed, ridges shorter than 1 mm.Ray flo rets white, female, 4 or 5, lamina shorter than 1 mm.narrower than 0.5 mm, tapering towards tip, 1-3-lobed: lobes equal or very uneven; tube 4.5-5.0mm long.D isc florets white; tube often tinged with pink below lobes, bisexual.5.0-5.5 mm long.5-lobed; lobes often asymmetrical.C ypselas subterete.± 1 mm long, greyish, inconspicuously ribbed, papillose, developing in 3-5 florets, more often in ray florets, released in a cluster at maturity, with pappus interlocking.Pappus bristles 18-23, ± 5 mm long, barbed in lower fifth, plumose in upper part, seta up to 1.5 mm long.
Flowering time: January-April.
Distribution and habitat: known from the Farm Klein Sleutelfontein in a dry riverbed in the Houtrivier, a trib utary of the Sand River, near Prince Albert and from an unknown locality of Drêge in the 'Bockland' (Figure 4).In deep shale-derived soils, in river or sand beds amongst grassy tufts.This species was recollected in September 1998 in the same area where Drêge originally found it.Although it appears to be quite rare, an extensive search in the area is needed to establish the extent of the distri bution.
Conservation status: rare, but well-conserved on pro tected farmland: could be endangered by flooding.long, foliaceous in upper part and along midvein, ovate, increasing in length from outer to inner series, outer about half the length of innermost bracts.Jan uary-M arch.
D iagnostic characters: compact grey bush; small, unobtrusive capitula; lamina of ray florets rolled like a watch-spring.

D istribution and habitat:
Western Cape between Nieuwoudtville and Clanwilliam (Figure 4).Sandy flats and mountain foothills.
The specific epithet 'grisea' refers to the blue-grey colour of these shrubs.

N u m b e r o f c o l l e c t i o n s k n o w n :
The specific epithet of this plant is to commemorate my father.Rudolph, in gratitude for his patient assistance in collecting specimens and seed of this species.
Wiry, robust shrub up to 1.2 m tall and 1 m wide.Branches smooth and very brittle, rarely more than 5 mm diam., very much entangled to form dense, sparselyleaved bushes; older branches bare with conspicuous leaf scars.Leaves very shortly petiolate, ascending, linear to narrowly oblong, twisted ± 270°-360°, greyish green, margins entire, slightly inrollcd.adaxial surface woolly, generally 6-10 x 1-2 mm, on new shoots to 30 x 5 mm, probably persistent for one season only.Brachyblasts generally absent, except on new, actively growing branches and then only 2 ^-le a v e d shoots.Capitula dis coid, homogamous, 7-10 mm long, 1-2 mm wide, con sistently 5-flowered; heads developing in the 2-7 termi nal leaf axils of a branch, linear when young, involucre spreading in upper half at maturity, lower part remaining tubular.Involucre with 30-35 bracts, bearing character istically tomentose markings on adaxial surfaces, fairly uniform in texture and appearance, becoming progres sively longer from outer to inner bracts.O uter involucral bracts scale-like, 1.5-2.0mm long and almost as wide, thus very widely ovate in shape, following series widely ovate, up to 2.5 mm long, next two series ovate, up to 4 mm long and 5 mm long respectively, innermost series lanceolate to linear, up to 8 mm long.Inner involucral bracts 10-15, in 4 or 5 series, scarious along margins, with multicellular glands, 6.0-7.5 mm long, linear to nar rowly oblong, those in outer series remaining green around midrib, hyaline towards apex, those in innermost series hyaline in upper half, mucronate, as long as pap pus.Receptacle flattened, less than 2 mm diam.. honey combed, ridges shorter than 1 mm.D isc florets white, bisexual.5-lobed.6 -7 mm long; lobes 0.8-1.0mm long.A nthers basally feathery-tailed.C ypselas subterete, asymmetric.± 1 mm long, greyish coloured, papillose, indistinctly ribbed, with 3 ribs visible, developing in only 1 of the 5 florets; ectocarp loose, easily removable, 1 cell layer thick; endocarp darker and thicker, forming a cavi ty filled with a hygroscopic gel-like substance, which expands rapidly when it comes into contact with water, causing the seedcoat to burst; cotyledons dark purplish; released in a cluster together with entangled pappus at maturity.Pappus bristles 20-28.± 5.5 mm long, barbed for a very short distance at base, plumose in upper part, seta up to 1 mm long.Flowering time: February-June.
D iagnostic characters: large bush; branches brittle; leaves widely spaced; capitula small, terminal on branch es; involucre spreading to release fluffy pappus; usually only one fairly large, subterete seed per capitulum.
The specific epithet of this plant is to commemorate my mother, Susan, who very enthusiastically helped to collect specimens at Kagga Kamma.
Wiry, erect, multistemmed, woody, yellowish grey shrub, up to 0.5 m tall.Stems up to 2 mm diam., bare in lower parts.Leaves 3 -7 mm long, sessile, mostly soli tary, but sometimes in small fascicles in young growth, ascending, linear, margins entire, slightly inrolled, adaxial surface woolly, leaf tips rounded; young leaves yel lowish green, older leaves fading to dark grey and drop ping off easily.Capitula discoid, homogamous, 6-7 x 2 mm, 5-flowered, terminal, solitary or in groups of up to 4, their bright, greenish yellow colour contrasting with the darker leaves.Involucre of 20-28 bracts, increasing in length from outer to inner series, 5.5-6.0 mm long, with a constriction at about one fifth from tip and yel lowish below constriction.O uter involucral bracts broadly ovate, with acute hyaline tips, not foliaceous, following series with hyaline part above constriction, gradually increasing in width and resulting in a membra nous, spoon-shaped tip in innermost bracts.Receptacle conical, less than 2 mm diam., honeycombed, ridges shorter than 1 mm.D isc florets white, bisexual.5-lobed, 4.5-5.0mm long, outer lobes of closed florets very glan dular.C ypselas subterete, ± 1 mm long, chestnutcoloured, inconspicuously ribbed, papillose, appearing to have a loose ectocarp, with a small attachment area between endo-and ectocarp at the base.Pappus bristles 20-26, ± 4.5 mm long, barbed in lower fifth, plumose in upper part, seta up to 1.5 mm long.Flowering time: November and December.D iagnostic characters: yellowish sheen to leaves and capitula; terminal heads small; involucre yellowish in lower parts with tips hyaline; innermost involucre with spoon-shaped tips.Erect, robust, single or multistemmed shrub, to 0.6 m tall, aromatic.Stems gnarled, dark grey, up to 40 mm diam., often buried in sand towards base; secondary branches up to 5 mm thick, bark thin, golden chestnutbrown to reddish brown or reddish in places, glabrous or with golden flakes pealing off; tertiary branches whitetom entose when young, becoming golden-yellow, glabrous and peeling when older.Leaves alternate, ses sile, linear, 1.0-11.0x 0.7-2.2mm, broadest at base, ascending but spreading in upper half, margins involute; upper surface canaliculate, w oolly-white; abaxially glabrous, epidermis sloughing off and becoming folliculate, especially at base; midrib raised, particularly at base, apex obtuse, mucro very small, inconspicuous.Capitula discoid, solitary, terminal, sessile, homoga mous, discoid, cylindrical, 7.0-8.0x 2.5-3.5 mm, 10-12-flowered.Involucral bracts 20-26, becoming progressively longer inwardly, tips hooded, pellucid, bronze, innermost bracts glabrous, lanceolate, up to 6.0 x 0.8-1.0mm, remaining bracts ovate, 1.5-6.0x 1.5-2.0mm, stereomes entire, with conspicuous white-tomentose patches at apices.Receptacle honeycombed, ridges between cypsela bases shorter than 1 mm.Erect woody, single-stemmed, brownish grey, strong ly aromatic, compact dwarf shrub, up to 0.2 m tall; very old plants up to 0.45 m tall but then breaking up.with main branches prostrate.Hark of older stems becoming greyish and flaky with age.tearing off in strips to uncov er a chestnut-coloured stem; older main stems up to 30 mm diam.. tending to split and appearing very uneven when seen in cross section, dead wood persistent on plant for a long time.Main branches up to 5 mm diam., prof usely branching into very fine shoots, less than 1 mm diam., with active growth restricted to very tips of fine branches, which are covered in minute leaves.Leaves up to 1.5 mm long, scale-like, adpressed, sessile, greyish green, alternate, about a leaf length apart, margins entire, slightly inrolled, adaxial surface woolly.Capitula dis coid.small, up to 7 x 2 mm.homogamous, 3-4-flow-ered.terminally borne on branchlets, linear when young, involucre spreading at maturity to release pappus and seeds.Outer involucral bracts 22-28, 1-6 mm long, ovate with acuminate tips, becoming progressively longer inwards, outermost scale-like.Inner involucral bracts linear, membranous in upper fifth, this part slight ly ascending, apex shortly mucronate.R eceptacle coni cal.less than 2 mm diam., honeycombed, ridges shorter than 1 mm.D isc florets white, bisexual, 5-lobed, 5.0-5.5 mm long.Cypselas subterete.± 1 mm long, greyish, inconspicuously ribbed, papillose, often only one per capitulum.appearing to have a loose ectocarp, but cavi ty with gel-like substance not obvious as in A. rudolphii.Pappus bristles 18-23, ± 5 mm long, barbed in lower fifth, plumose in upper part, seta up to 1.5 mm long.Flowering time: January-M arch.
D iagnostic characters: compact shrub with branches copiously branched into very fine, secondary branches; leaves and twigs pleasantly aromatic when crushed.
Distribution and habitat: restricted to Kovis M oun tain near Luderitz in Namibia, possibly also on other mountains in the area (Figure 7).In sand on granite mountains.
Conservation status: not threatened.Woody, compact, multistemmed dwarf shrubs, up to 0.6 m tall, but very often with growth stunted and then ± 0.2 m tall.Stems up to 3 mm diam., smooth, wiry and brittle, produced from a well-developed rootstock, older dead stems persistent; stems bare for largest part, leaf cluster scars prominent, profusely and very finely branched in upper parts, young growth often with zigzag appearance.Leaves sessile, in fascicles; subtending leaves up to 15 x 1 mm.about twice as long as others in fascicle, ascending, linear, straight or twisted, margins entire, slightly inrolled, adaxial surface woolly; fascicles widely spaced, at least a leaf length apart, leaving

D
istribution and habitat: on plateau on Swartruggens Mountain in the Ceres-Karoo (Figure7).Deep red or white sand in level areas between TMS outcrops.Conservation status: not threatened.

D isc florets FIGURE 7 -Geographical distribution of A susannae, • . and A thuja, ♦
D

istribution and habitat: it
has a restricted distribu tion in the northwestern area of the Witsand dune com plex, 70 km southwest of Postmasburg in the Northern Cape (Figure6).White sand dunes of Aeolian origin, deposited during the Middle and Upper Pleistocene, as were the red Kalahari sands.The water table in the dune complex is high and acts as a reservoir to support vege tation in the basin.status: not threatened.The Witsand area is now being conserved by Cape Nature Conservation and as long as the habitat is protected, there is no threat of extinction.Although it is localised, it is abundant in the area.