Revision of Melhania in southern Africa

The 13 species o f Melhania in southern Africa are revised. Among the problems resolved is a long-standing one concerning the identities o f M. rehmannii Szyszyl. and M. griquensis H. Bol., the latter in its strict sense being relegated to synonymy o f the former.

Shrublet, 20-60 cm tall, sometimes up to 1 m tall, branched from near the base, erect or, when browsed, forming a low bushy plant, main stem woody; branchlets tomentose, the grey tomentum in termixed with light brown, clustered, villose hairs.Stipules subulate, tomentose, up to 14 cm long.Leaves pubescent above with subappressed, long, single hairs or very occasionally 2 or 3 hairs from the same base, greyish-stellate-tomentose beneath inter mixed with clustered light brown hairs, especially on the nerves, ovate-oblong to ovate-lanceolate, up to 10 cm long and 4 cm broad, margins shallowly den tate, sometimes obscurely so; petiole 0,7-1,5 cm long.Inflorescence axillary, peduncle rigid, up to 5 cm long, usually 2-flowered at the apex, pedicels up to 1 cm long.Epicalyx-bracts, broadly ovate, acumi nate, cuspidate at the apex, cordate at the base, usually slightly longer than the calyx and petals, According to Harvey in F. C. 1: 222, M. leucantha E. Mey.nom.nud. is this species.The Drege speci mens at Kew which Meyer named M. leucantha have been examined and they are M. didyma.
M. didyma resembles M. forbesii in some respects and especially in the size and shape of the epicalyxbracts, but it is readily distinguished by the upper surface of the leaf being pubescent with single subappressed hairs, while in M. forbesii it is tomentose or with short, grouped or stellate hairs.
The specimens from Botswana listed by O. B. Mil ler in J 1 S. Afr.Bot. 18: 57 (1952)  Shrublet, a weak to fairly strong perennial, usually branching near the base; branches upright, spreading or prostrate, usually branched again 20-60 cm long, new growth grey-stellate-tomentose, tomentum inter spersed in parts with scattered red-brown, bunched hairs.Stipules subulate 4-14 mm long.Leaves linearlanceolate linear-oblong, narrowly oblong or nar rowly ovate-oblong, rounded at base and apex, rarely apex truncate and very shallowly lobed, 2,4-12 cm long, 0,4-2 cm broad, rarely 2,5 cm broad, discolourous, lower surface grey-tomentose with golden-brown, lepidote, stellate-pilose scales bearing short or long hairs upper surface fairly sparsely pubescent with mostly rather long single, (or rarely about 3 in a cluster), subappressed hairs, lying i in the same direction or glabrous; margins entire, very rarely shallowly lobed in part; lateral veins few, im pressed above, prominent beneath, not excurrent; petiole 0,2-1,2 cm long, rarely 1,5 cm long, grey tomentose.Inflorescence axillary 1-2-flowered; peduncles 3-32 mm long, ultimately longer than the petioles which range from 2-12 mm, frequently in the single flowered inflorescence the peduncle and pedicels are indistinguishable.Epicalyx-bracts from narrowly to broadly ovate-acuminate, shallowly to deeply cordate at the base and abruptly narrowed in a short basal claw, about 8-15 mm long and 2-6,5 mm broad, thinly to densely stellate-tomentose on both surfaces.Calyx with lobes ovate-lanceolate, acumi nate, shorter or longer than the epicalyx, 8-14 mm long, 2,5-4 mm broad, dorsally grey-tomentose, sparsely to very densely interspersed with bunched or stellate golden-brown, bristle-like hairs.Petals yellow, broadest at the apex cuneate, about 12 mm long and 8 mm broad at apex.Stamens about 4 mm long, alternating with linear-ligulate staminodes somewhat longer than the stamens.Ovary subglobose densely tomentose and bristly with bunched or stellate hairs; style 1,5-3,75 mm long with branches half to three quarters as long; ovules 6-8 in a cell.Fig. 2. The decision to follow the authors of Flora Zambesiaca and put M. linearifolia Sond.into synonymy under M. prostrata and not as a variety of M. didyma as Szyszylowicz did, was made after seeing the type specimen of the former, Gueinzius 532.This speci men compares well with the type of M. prostrata from Griqualand West and the many specimens of this species found throughout the Transvaal and in parts of Natal.
M. prostrata is closely related to A/, didyma, but may be distinguished by the much narrower leaves with entire margins (very rarely a single leaf shallow ly lobed in part).The pubescence on the upper sur face of the leaves is, when present, like that of M. didyma of rather long usual single subappressed hairs, but in M. prostrata the upper surface is early glabrescent.
Where the distribution of these two species overlap, intermediates are found.It is not clear whether these intermediates are of hybrid origin or whether the two taxa are fairly recent segregates from a common stock.
The specimen of Schlechter 4322 in the Transvaal Museum Herbarium, now in the National Herb arium, is a mixture of M. prostrata and M. transvaalensis.The latter has the upper surface of the leaves distinctly stellate throughout.
When Burchell collected the type of this species, Burchell 2153, he evidently thought it was a Dombeya, for in his catalogue he has listed it as Dombeya prostrata.
Shrublet, 20-45 cm tall with several slender woody stems from a woody rootstock, laxly branch ed; branches shortly and densely tomentose, some times, in new growth reddish-brown lepidote stellate scales are obvious.Stipules subulate, 6-15 mm long.Leaves on flowering branches clustered towards the apex, entire (very rarely lower leaves shallowly tooth ed in part), finely and densely velvety tomentose on the upper surface, greyish tomentose beneath with reddish, lepidote, stellate scales usually obvious, es pecially on the nerves, narrowly lanceolate-elliptic, mostly 2-6,5 cm long and 0,4-1,2 cm broad, sometimes up to 1,8 cm broad, nerves prominent beneath, obscure above, basal nerves 3, rarely 5, base subrounded, apex subacute or rounded, mucronate; petiole 0,4-1 cm long, tomentose with a few to many reddish-brown, lepidote stellate scales obvious.In florescence axillary in the upper leaves, 1-3-flowered; peduncle 4-20 mm long, pedicels 3-14 mm long, both tomentose with scattered reddish-brown stellate scales obvious in parts.Epicalyx-bracts ovate acuminate, rounded to cordate at the base, about 12 mm long, 6 mm broad near the base, tomentose on both surfaces with the lepidote scales obvious on the outside.Calyx-lobes lanceolate, acuminate, about 15 mm long, 3,5 mm broad near the base, tomentose and lepidote without.Petals canary yellow, turning brownish at maturity, broadest at the apex, about 16 mm long, 10-14 mm broad at apex.Stamens with filaments 1 mm long, anthers 3-4 mm long, staminodes 9 mm long.Ovary subglobose, about 3 mm diam., white tomentose; style about 6-8 mm long, branches 1,5-3 mm long; ovules 7 in each locule.Capsule about 10 mm long, 8 mm diam., shortly tomentose and scaly, the scales bearing short hairs.Fig. 3. Recorded from the Wolkberg, the northern ex tremity of the Transvaal Drakensberg and at the top of Abel Erasmus Pass, growing on rocky hillsides among grass in shallow soil over dolomite.This species occurs abundantly in the area which lies south-east of Boyne in the Pietersburg District and on the borders of the Letaba District.It grows on the grassy slopes among dolomite rocks.The entire, velvety-tomentose leaves, together with the long style and large petals, characterize the species.
It resembles M. randii Bak.f. from Zimbabwe in the entire leaves but differs, among other things, in the pubescence on the upper surface of the leaves, the larger flowers and the ovate-cordate epicalyx.The specimen from Abel Erasmus Pass is not quite as velvety as the Wolkberg specimens.
Shrublet, 6-60 cm tall with several slender woody stems from a thick, woody rootstock, branches at first finely stellate-tomentose with minute reddish brown scales intermingled.Stipules narrowly lanceolate-subulate up to 8 mm long.Leaves entire, rarely some shallowly dentate in part, finely minutely stellate-tomentose and with scattered reddish brown scales (the scales sometimes stalked and all bearing short hairs) on the under surface, the upper surface finely stellate-tomentose and in some forms early glabrescent, narrowly oblong-elliptic or lanceolateelliptic to ovate-oblong, 2-9 cm long, 0,2-1,5 cm broad, rounded at the base, subacute to rounded at the apex, mucronate; petiole up to 1 cm long, tomen tose and lepidote.Inflorescence axillary, tomentose and lepidote, 1-3-flowered; peduncle 7 mm long and indistinguishable from the pedicel or up to 4 cm long and pedicels up to 1,2 cm long.Epicalyx-bracts nar rowly lanceolate, narrowly ovate to ovate-acuminate, from 6-13 mm long and 1-6 mm broad (large bracts found mostly on the specimens from the eastern mountainous country), tomentose on both surfaces and with minute reddish scales intermixed on upper surface.Calyx-lobes lanceolate-acuminate, up to 11 mm long, tomentose and with conspicuous reddish brown scales on outer surface.Petals yellow, 'bright golden yellow' about 9 mm long, 7 mm broad at the oblique apex.Stamens with filaments 1 mm long, an thers 2-3 mm long, staminodes 3-5 mm long.Ovary stellate tomentose and with straight hairs especially in upper portion, about 2 mm diam., style short, at maturity up to 2 mm long, branches when spreading up to 3 mm long.Capsule up to 1 cm long, stellate-pubescent and scaly.
Recorded from the Middelburg District, where it grows in open mixed grassveld in iron-rich soil and a taller form from the Barberton District on the expos ed, wind-swept summit of a mountain near Barber ton.Also occurs in Zimbabwe and Mozambique.M. randii has been recorded from four widely separated areas of distribution.Probably the long period of isolation accounts for the plants in any one of these colonies differing in some respects from those in another.The specimens seen to date from around Salisbury, the type locality; are low bushes with long narrow leaves on which the pubescence per sists on the upper surface.In the mountainous eastern regions of Zimbabwe and the Transvaal the plants are over 30 cm tall with slender erect stems; they also have long leaves with persistent pubescence on the upper surface, and differ only slightly from each other, for instance the epicalyx bracts on the Transvaal specimens are usually broader than those on the Zimbabwean specimens.The form found in the Middelburg District of the Transvaal differs from all the others in having shorter and broader leaves, which are early glabrescent on the upper surface.In this form the bushes are usually under 30 cm tall.
M. randii resembles M. prostrata and M. Integra in the entire leaves.It can be distinguished from the former, among other things, by the upper surface of the leaves being finely stellate-pubescent before becoming glabrous as opposed to having long simple, appressed hairs on the upper surface, before becom ing glabrous.From M. Integra it may be distinguish ed by the persistent minutely stellate pubescence of the upper surface of the leaf as well as the short style and smaller flowers, the styles being 1,75 mm long as opposed to 8 mm long in M. Integra and the flowers 1,1 cm long as opposed to being 2 cm long.Shrublet about 60 cm tall, sometimes taller, stem erect, branched; branches suberect the upper portion usually appearing rust-coloured from the clusters qf rather long ferruginous hairs, which are sometimes stalked and occur more or less densely intermingled with the short greyish stellate tomentum.Stipules subulate 10-20 mm long.Leaves greyish tomentose with short, fine grouped or stellate hairs especially dense on the lower surface, and sometimes with clusters of ferruginous hairs on the nerves beneath, more or less ovate-oblong, 3-11 cm long, 1, 5-6 cm broad, rounded at the apex, sometimes mucronate, base rounded to cordate, margin crenate-serrulate; petiole 1-2,5 cm long, usually densely covered with clusters of ferruginous hairs.Inflorescence axillary, peduncles straight, tomentose and with numerous clusters of ferruginous hairs, suberect, 1-6 cm long, branched near the apex; raceme 1-4-flowered, usual ly 3-flowered; pedicels 2-7 mm long.Epicalyx-bracts from about 12-18 mm long and 10-14 mm broad, ovate, sometimes broadly acuminate to an acute apex (not abruptly so), cordate at the base, longer than the calyx and petals (in dried specimens obscuring the calyx), tomentose on both surfaces.Calyx-lobes greyish villose-tomentose dorsally, glabrous within, lanceolate about 1 cm long, apex acute.Petals about as long as the calyx-lobes, obovate, broadest at the apex (the faded petals twisted at the apex and per sisting like a cap on the capsule may appear to be longer than the calyx but they are not attached at the base).Stamens about 12 mm long, filaments and an thers about 5 mm long.Ovary subglobose about 9 mm diam., densely tomentose; style about 2 mm long, style-branches about 4 mm long.Seeds up to 6 in a cell.

Melhania forbesii
Found along rivers in alluvial soil or on dolomitic hillsides, sandy flats and parkland in red soil.Recorded from Natal, Swaziland, the eastern and northern Transvaal; and northern South West Africa.Also occurs in Mozambique, Zambia, Zim babwe and Angola.In the shape of the epicalyx M. forbesii closely resembles M. didyma, but can be readily distinguish ed by the upper surface of the leaves being tomentose and minutely stellate and not pubescent with single, long, sub-appressed hairs as in M. didyma.The specimens from Botswana listed by O. B. Miller in J1 S. Afr. Bot. 18: 57 (1952) as M. didyma are M. forbesii and, most probably, Dinter 3036 also listed as M. didyma from Tsumeb, in Feddes Reprium 19: 96 (1924), is this species.(See also notes under M. acuminata var.acuminata.) The native name for M. forbesii is moulhwadambo meaning 'setting sun'.
Shrublet about 35 cm tall, with a woody rootstock.Stems many, rather robust, in dried specimens 3-5 mm diam., laxly branched, new growth with a sul phur-grey stellate tomentum, the hairs of different lengths and grouped on scales.Stipules subulate, 5-10 mm long.Leaves finely and densely sulphurgrey stellate-tomentose on both surfaces, upper sur face somewhat darker than the lower; lamina broadly ovate-trullate or ovate-elliptic, 3-9, 5 cm long, 1,5-5,5 cm broad, broadly cuneate or subrounded at the base, narrowing slightly to the obtuse or subtruncate apex, shallowly crenate-dentate except at the base; petiole 1-2 cm long, densely sulphur-grey tomentose, fairly stout.Inflorescence in axils of up per leaves; peduncle up to 6 cm long, stellatetomentose, 1-3-flowered at apex; pedicels 0,3-1,5 cm long.Flowers polygamous.Epicalyx-bracts ovate, cordate at base, gradually attenuating to a subulate upper portion and acute apex, longer than calyx, about 14 mm long, 4-12 mm wide, stellatetomentose on both surfaces.Calyx with lobes deltoid-acuminate, about 11 mm long, outer surface stellate-tomentose.Petals yellow, broadest at the apex, in essentially male flowers about 14 mm long, in female shorter, about 10 mm long.Stamens, in male flowers, with anthers about 4 mm long, in female abortive, about 2 mm long; staminodes about 6,5 mm long.Ovary densely beset with long, grouped hairs on scales; style in male flowers long and slender, about 7 mm long, branches aborted or short, style in female flowers thick and short, about 2,5 mm long, branches longer than the style, about 5 mm long, sometimes decurrent on the style; ovules 5 in a cell.Capsule broadly oblong, about 9 mm long, tomentose and with groups of rather long, sulphur-coloured hairs.Fig. 4.
To date recorded only from the Umfolozi Game Reserve, Zululand, where it grows on grassy hillslopes.
N a ta l .-LowerUmfolozi: Umfolozi Game Reserve, Ward 3324; Dengeseni Beacon Hill, Strey 4957a; 4957b; 4957c; 4957d.This is the first known record of a species of Melhania with polygamous flowers.The essentially male flowers are more showy with longer petals, large stamens and an aborted style.The female flowers have short petals, small stamens and a style with long, stout branches.The broad ovate-trullate leaves, that is, leaves, which are usually broadest about a third of length above the base, are a distin guishing feature.Shrublet about 65 cm tall; stem erect from a com paratively slender tap-root, branched; branches erect and spreading, new growth greyish stellate-tomen tose and sometimes with bunched light to dark brown hairs intermixed.Stipules subulate, 0,5-1,7 cm long.Leaves greyish stellate-tomentose on both surfaces, sometimes more thinly so on the upper surface, lower EE3 surface sometimes with light to dark brown bunched hairs, especially on the prominent nerves, broadly to narrowly ovate-oblong, 3,5-10 cm long, 1,5-6 cm broad, narrowing towards a broadly rounded apex, rounded to subcordate at the base, margins crenateserrate; petiole 1-2,5 cm long, stellate-tomentose.Inflorescence axillary, peduncle straight, suberect up to 5 cm long, 3-or more-flowered near the apex; pedicels 0,6-1 cm long.Epicalyx-bracts ovate (rarely narrowly so), shorter than the calyx, rounded at the base and with a very short broad claw, abruptly acuminate above into a caudate-like upper half, tomentose on both surfaces.Calyx with lobes lanceolate, gradually narrowing to the acute apex, 1-1,5 cm long, stellate-tomentose on the outside on ly.Petals usually longer than the epicalyx, obovate, broadest at apex.1-1,5 cm long.Stamens with filaments and anthers about 3,5 mm long; staminodes 7 mm long.Ovary subglobose about 8 mm diam., densely tomentose; style 6-10 mm long, branches 1-2 mm long; ovules 6 in a cell.
For key to varieties see key to species.The typical variety is characterized by the leaves usually being broadly ovate to ovate-oblong, the flower large, and the rather thick tomentum being predominantly grey with fairly obscure, bunched, brown hairs intermixed.There is a fair number of herbarium specimens, which approach var.agnosta either in the leaves being narrower than usual or the bunched hairs somewhat more conspicuous.This supports the decision to reduce M. agnosta K. Schum. to varietal rank under M. acuminata.
Found in sandy soil, on flats, in bushveld and on grassy wooded slopes.Recorded from the northern Cape, the northern and eastern Transvaal, South West Africa and Botswana.Also occurs in Zambia, Zimbabwe, Angola and Mozambique.A/, acuminata var.acuminata is sometimes con fused with M. forbesii, which has more or less the same habit and partly the same distribution.These taxa are mainly distinguished by the shape and length of the epicalyx-bracts around the length of the style.
According This variety differs from the typical in the leaves being narrowly oblong or narrowly ovate-oblong and the tomentum on the newf growth being densely inter spersed with bunched, dark-brown hairs.The flowers are smaller on the whole and usually clustered at the apices of the branchlets.It agrees with the typical in the characteristic abruptly-acuminate epicalyxbracts, the long style, general habit and the long peduncles bearing 1 to 4 flowers at the apex.
Recorded from Botswana and the Transvaal.Also occurs in Zambia, Zimbabwe and Mozambique.A specimen collected in 1978 on the Pilanesberg, Rustenburg District, Peeters, Gericke & Burelli 590 may be this variety although the leaves are somewhat broader than in the type specimen.
Found on stony hills and grassy slopes on dolomite formation.Recorded from restricted areas in the nor thern Cape and the Transvaal.A study of Rehmann's route when he collected the type of this species shows that the locality 'Elandsriver Drift' was most probably in the northern part of the Pretoria District, north-east of Rust de Winter.
The herbarium sheet of Schlechter 4322 from the Transvaal Museum, now in the National Herbarium, Pretoria, has mounted on it a mixture of this species and M. prostrata.Another sheet of the same number, Schlechter 4322, which has been in the Na tional Herbarium for many years, is purely M. transvaalensis, i.e. all the leaves are crenate-dentate and coarsely stellate on the upper surface (not entire with simple subappressed, long hairs or glabrous on the upper surface).
Found in bushveld, sandy flats or rocky ridges in dry country.Recorded from the north-western Cape, Swaziland, the central and northern Transvaal, Bot As explained by Wild in Flora Zambesiaca, the confusion between M. rehmannii and M. griquensis (now' pro parte under M. virescens) was initiated by H. Bolus, the author of M. griquensis who, although basing his description on the Orpen specimen from Griquatown (hence griquensis), cited Burchell 2050 from the Asbestos Mountains as the holotype: he saw this specimen at Kew in 1881 and mistakenly con sidered it to be conspecific with the Orpen specimen.Subsequent taxonomists such as K. Schuman, Wild and Burtt Davy all agreed that two taxa were invol ved, for although closely related, they were readily distinguished by the epicalyx bracts which were subulate to linear in the Griquatown specimen and broadly ovate in Burchell's specimen from Asbestos Mountains.Following these taxonomists, it has become necessary to relegate M. griquensis sensu stricto to synonomy of M. rehmannii, and M. gri quensis (two parte as to the description and Orpen specimen) to M. virescens (K.Schum.)K. Schum., the first published synonym.
For further distinguishing features and notes on distribution see under M. virescens.
Found in limy soils in the northern Cape, the Transvaal, Botswana and South West Africa.The confusion that existed between this species and M. rehmannii is understandable for, until one has recognized the features that distinguish them, these species look very much alike.The specimen on which Bolus based the greater part of his description of M. griquensis, i.e.Orpen sub Bolus 6015, is a mixed gathering.Four of the portions have the linearsubulate epicalyx bracts as described by Bolus, and the fifth, on the left hand side of the sheet, the ovate bracts of M. rehmannii.the distribution of these two species overlap in the northern Cape and parts of the Transvaal.A further distinguishing feature is that the peduncles in M. virescens are short (usually shorter than the petiole) and suberect and the pedicels, if pre sent, are also short and suberect, whole in M. rehmannii the peduncles are somewhat longer (about as long as or longer than the petiole) and the pedicels, if present, longer, thinner and cernuous.
With regard to the distribution, as mentioned above, these species overlap in the northern Cape and the Transvaal, but M. virescens extends further westward in South West Africa.According to collec tors' notes M. virescens is restricted to limy soil, while M. rehmannii is not so restricted and is much more widespread.
The specimens added by N. E. Brown to Bolus's original description of M. griquensis, namely Bur chell 2385, Holub s.n.Rehmann 5220 are all M. rehmannii.
Found in hot, dry parts of the north-western Cape, the Transvaal, Orange Free State, South West Africa and Botswana.Also occurs in Zimbabwe.In the Transvaal this species is very common be tween Potgietersrus and Pietersburg.The form which occurs here agrees with the description of M. albi cans, but no grounds can be found for separating it from M. burchellii in spite of the disjunct distribu tion and the difference in habitat.In the Transvaal it grows on a heavy gravelly loam soil which is very dif ferent from the loose sand in which it is found in the north-western Cape.It may be noted that the type locality of M. albicans is given as Pilgrims Rest.The Rev. William Greenstock, who collected the original specimen, spent some time with Edward Button at Eersteling Mine, between Potgietersrus and Pieters burg, before proceeding to Pilgrims Rest.The plant has not been found at Pilgrims Rest again and it is considered that Greenstock collected this specimen in the neighbourhood of Eersteling.Several similar cases are known where species based on Greenstock specimens have been attributed to Pilgrims Rest, but subsequent investigation has shown that they were without doubt collected in the Eersteling area.
In the Flora Capensis under M. burchellii Miss Owen's specimen is cited as coming from 'Zooloo Country'; this is probably another case of the several instances where Miss Owen's specimens, collected after she had left Zululand (that is, after the Dingaan massacre), were labelled as from Zululand, whereas they probably were from the Kuruman or Zeerust Districts.
Regarding another citation in the Flora Capensis, namely Zeyher from the 'Aapies River', it is doubtful whether this locality is correctly cited, since this species has not been found near Pretoria.
Although none of the syntypes of M. dinteri has been seen, the description best fits the specimens in this taxon.
Occurs throughout the length of western South West Africa with one record to date from the nor thern Cape.Found mostly in dry conditions on the western side of the plateau at the Namib fringe.This species approaches M. suluensis from eastern Natal.For main distinguishing features see the key, p. 264.
M. damarana also resembles forms of M. ovata auct.found in tropical Africa with the type from South America, the principal distinguishing feature being the shape of the epicalyx-bracts, which are linear-subulate in M. ovata and lanceolate to broadly lanceolate in M. damarana.
Found in alluvial soil or on rocky slopes in dry bushveld at altitudes of 3-170 m.Recorded from the coastal area of Natal.M. suluensis is closely related to M. damarana, which occurs in South West Africa, west of the escarpment.Besides this difference in distribution the main distinguishing features are given in the key on p. 264.

ACKNOWLEDGEMENTS
When reviewing the genus Melhania in 1962, I was fortunate in being able to enlist the help of Dr O.A. Leistner, who was the South African Liaison Officer at the Royal Botanic Gardens, Kew: he kindly ex amined specimens for me at Kew and the British Museum (Natural History) and solved the problems submitted to him.I am indeed grateful to Dr Leistner for his assistance.
1 am also indebted to Prof. H. Wild of the Univer sity of Zimbabwe and Mr R. B. Drummond of the Salisbury Herbarium, Zimbabwe, for advice and the loan of specimens of Melhania randii.
The loan of herbarium material from Rhodes University and the University of the Orange Free State was much appreciated.

Blue Jay Ranch, Com pton 29746.
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