Four new species of Erica (Ericaceae) from Western Cape, South Africa

Four new species of Erica L. from the mountains of Western Cape are described: E. richardii E.G.H.Oliv., rare and localized on quart/ite cliffs in the foothills of the Great Swartberg north of Klaarstroom, E. anemodes E.G.H.Oliv., and E. viminalis E.G.H.Oliv., both restricted to the Hex River and adjacent mountains, and E. limnophila E.G.H.Oliv., rare around high altitude marshes in the Wemmershoek and Dutoitskloof Mountains. 1. Erica richardii E.G.H.Oliv., sp. nov., E. flocciflorae Benth. ct E. saxigenae Dulfer affinis propter formam seminum sed a prima floribus albis corolla puberula, habitu tenello non crasso lignoso et a secunda floribus albis corolla minore et habitu tenello non crasso lignoso differt. Figura 1. TYPE.—Western Cape, 3322 (Oudtshoom): Klaar­ stroom area. Witberg southeast of Farm Droekloof, north­ ern slopes near summit. 1 140 m, (-BC), 12 July 2000, Oliver 11529 (NBG. holo.; BM, K, MO, NY, PRE, S). Shrubs varying from very small, ± 10 x 50 mm and prostrate, to erect and divaricate, up to 250 x 3(X) mm, woody, single-stemmed reseeder. Branches: few to many main branches (10-)20-30(-60) mm long terminating in a florescence, no secondary branches, with ± 3 new branch­ es formed each year just below each florescence, all cov­ ered with dense short retrorse white hairs. Leaves 3-nate, erect, imbricate, linear, 3.5(-4.8) x 0.8 mm, adaxially slightly rounded, abaxially rounded, both sides with very short scattered hairs becoming glabrous, margins subacute with non-sticky glands in young stages, apex acute some­ times reddish, sulcus narrow open at base; petiole ± 0.6 mm long, glabrous, ciliate. Inflorescence: flowers 3-nate in 1 w horl at apex of main branches; pedicel ± 7 mm long, red, covered with dense short white hairs, these somewhat clustered together in stellate-like groups with few, nonsticky, very short-stalked glands; bract partially recaulescent 'A -1/; way up pedicel, oblong to oblanceolate, ± 2.4 x 0.8 mm, covered with short scattered hairs and few, short, non-sticky glands on margins, creamish white with red­ dish apex, sulcus narrow ± '/2 as long as bract; bracteoles 2. placed a short distance above bract, otherwise like bract. Calyx 4-partite; lobes not laterally imbricate, elliptic to obovate, ± 2.5 x 1.5 mm long, flattened base, white to creamish, with semitransparent margins, with stouter cream-coloured, sometimes reddish, sulcate upper portion, sparsely and shortly hairy, margins minutely serrated with occasional short, non-sticky, gland-tipped hairs, sulcus narrow ± 'A length of sepal. Corolla 4-lobcd. globose urceolate, ± 4.5 x 3.8 mm, covered with very short hairs, white; lobes ± 1 x 1 mm, rounded entire. Stamens 8, free. * Compton Herbarium, National Botanical Institute, Private Bag X7, 7735 Claremont, Cape Town. MS. received: 2(XM)-(W-12. included; filaments oblong broad, ± 2.0 x 0.3 mm, with distinct apical S-bend, glabrous, white; anthers bipartite, basally attached, lanceolate in adaxial view, appendiculate; thecae erect, ± 1.1 x 0.6 mm and ovate in side view, with long pale red hairs between thecae otherwise glabrous, red-brown with darker apex, appendages obo­ vate, ± 0.4 x 0.2 mm, pink-white, edged with long white hairs; pollen as tetrads. Ovary' 4-locular, globose, ± 1.8 x 2.0 mm, slightly tapered at apex, glabrous, green, soon turning purplish with well-developed dark red nectaries around base; ovules ± 20 per locule, spreading laterally from centrally placed placenta; style ± 2 mm long, mani­ fest, becoming exserted in fruiting stage, green with pur­ ple apex, glabrous; stigma truncate. Fruit a dehiscent loculicidal capsule, ± 3.5 x 3.5 mm, very hard and woody; valves splitting halfway open but suberect and spreading very little, septa mainly on the valves very thin and papery portion on columella. Seeds asymmetrically angularobovoid with folds and ridges, ± 0.9 x 0.6 mm, yellow occasionally tinged red, testa thickened, deeply reticulate, cells 50-75 x 40-60 pm, anticlinal walls straight abaxial­ ly, undulate adaxially, inner periclinal walls with nume­ rous small pits. Figure 1. This new species is related to E. flocciflora and E. sax­ igena, and to a lesser extent E. affinis Benth., mainly because of the very similar seeds, which are unusual in the genus with their asymmetrical shape covered with irregular ridges and folds. The capsules are also similar in being very woody and not opening very much. Other shared characters are the long hairy pedicels with coloured bract and bracteoles, the glabrous, globose ovary with a short style and truncate stigma, and the anthers with broad appendages and filaments. It seems to be closest to E. flocci flora and E. saxigena because all three possess hairy corollas and sepals, and have similar leaves. E. flocciflora differs in having the hairs noticeably long and woolly on the pedicel, bract, bracteoles and sepals, the flower colour being greenish yellow, the appendages broader and serrate (not ciliate) and in the shrubs being extremely woody and erect up to 1 m tall. E. saxigena differs in having no hairs on the anthers, erose appendages, a velvety, hairy, much larger crimson corol­ la and a stout woody growth. Erica affinis can be distinguished by the glabrous bright pink corolla and sepals, the latter being broad and 2 Bothalia 31,1 (2001)

This new species is related to E. flocciflora and E. sax igena, and to a lesser extent E. affinis Benth., mainly because of the very similar seeds, which are unusual in the genus with their asymmetrical shape covered with irregular ridges and folds. The capsules are also similar in being very woody and not opening very much. Other shared characters are the long hairy pedicels with coloured bract and bracteoles, the glabrous, globose ovary with a short style and truncate stigma, and the anthers with broad appendages and filaments. It seems to be closest to E. flocci flora and E. saxigena because all three possess hairy corollas and sepals, and have similar leaves. E. flocciflora differs in having the hairs noticeably long and woolly on the pedicel, bract, bracteoles and sepals, the flower colour being greenish yellow, the appendages broader and serrate (not ciliate) and in the shrubs being extremely woody and erect up to 1 m tall. E. saxigena differs in having no hairs on the anthers, erose appendages, a velvety, hairy, much larger crimson corol la and a stout woody growth. petaloid and as long as the corolla, and by the broader bract and bracteoles which are adpressed to the calyx. It also produces much larger shrubs up to 1.5, rarely 2 m tall.
Superficially the new species looks like the smallflowered forms of E. spectabilis Klotzsch ex Benth., E. eustacei L.Bolus, E. tragulifera Salisb. and E. formosa Thunb. in the shape and colour of the flower, the broad filaments, hairs on the anthers, and the long hairy pedi cel with similar bract and bracteoles. The seeds of these species, however, are quite different from those of E. richardii, being rounded and symmetrically shaped and simply reticulate. All four of these species occur in the mountains in and around the Little Karoo to the south and west of Klaarstroom.
Erica richardii has only recently been discovered in a habitat that is most unlikely for Ericaceae. It grows only on the north side of cliffs, boulders and rocky outcrops of quartzites of the Witteberg Series on the boundary of the dry arid Great Karoo ( Figure 2). This habitat occurs in the low mountain ridges north of the Great Swartberg  This species was sent to us in May 2000 by Richard Taylor of the Technicon at Saasveld, George, a keen climber and supporter of the Protea Atlas Project, who was trying to locate the most northern outliers of Proteaceae in the low foothills north of the Great Swartberg Range. He led us to the locality where we managed to locate 25 plants spread for some 4 km along the north-facing cliffs and upper part of the summit ridge. The plants were still in flower in early July with older fruiting flowers and some buds also present on some plants.
The plants of E. richardii grow only in crevices in the rocks and vary considerably in size. The younger plants and those in very small rock crevices on vertical faces are small and prostrate, whereas the older ones that have managed to grow in larger rock crevices or on more hor izontal rock faces, are divaricate-the largest one 300 mm tall. Most seem to have a very woody basal stem, which would indicate that they are all of considerable age. However, the branches are surprisingly rather thin and delicate with leaves confined to the upper parts, which is in stark contrast to the extremely woody, stoutly branched and leafy shrubs of E. flocciflora and E. saxigena.   es 2-10 mm long ending in an inflorescence; stems cov ered with dense short retrorse simple hairs, no infrafoliar ridges. Leaves 3-nate, slightly imbricate, erect to sub spreading, elliptic-oblong, ± 4.2 x 1.0 mm, adaxially flattened, abaxially rounded margins subacute, glabrous, minutely ciliate, when young with few, small, non-sticky glands on margins, narrowly sulcate, sulcus open at base; petiole ± 0.7 mm long, adpressed, adaxially puberulous, minutely ciliate. Inflorescence: 3 flowers in 1(2) whorls at ends of main and secondary branches; pedicel 3 mm long, curved, with dense, short, retrorse, simple hairs, occasionally a few plumose hairs intermixed; bract par tially recaulescent in middle of pedicel, ovate to elliptic, ± 2 x 1 mm, acute, glabrous, margins with small nonsticky glands and basally short hairs, white, sulcus nar row in upper V4; bracteoles 2, just above bract, elliptic to obovate, ± 1.5 x 0.9 mm, obtuse to subacute, otherwise like bract. Calyx 4-partite; lobes ovate to elliptic, ± 2.0 x 1.3 mm, laterally imbricate, adpressed to corolla, apex subacute and cucullate, white, glabrous, margins with small, sessile, non-sticky glands and basally few short hairs, sulcus xU -h length of sepal. Corolla 4-lobed, broadly campanulate, 2.5 x 2.5 mm, glabrous, white; lobes erect to spreading, ± length of corolla, ± 1.3 x 1.3 mm, triangular subacute, entire. Stamens 8, free, included to manifest; filaments ± 2 mm long, linearoblong, with strong apical S-bend, glabrous, white; anthers dorsifixed near base, bilobed, quadrate in adaxi al view, appendiculate; thecae erect, adpressed, ± 1.1 x 0.6 mm, ellipsoid in lateral view, dark brown, aculeate on inner and adaxial margins, appendages pendulous, lance olate, ± 0.5 x 0.2 mm, irregularly serrate and with few hairs, brown; pore ± x!z length of theca; pollen in tetrads. Ovary 4-locular, broadly ovoid to ellipsoid, 1.0 x 1.5 mm, obtuse, glabrous, with reduced nectaries around base; ovules ± 12 per locule, subpendulous from large placenta in upper half; style exserted, ± 2.3 mm long; stigma capitate-peltate, ± 0.7 mm across. Fruit a dehis cent capsule, broadly ovoid, ± 1.1 x 1.8 mm, valves split ting almost to the base but only slightly spreading, sep tum ± 90% on the woody columella and 10% on valve. Seeds ovoid-ellipsoid, ± 0.7 x 0.5 mm, occasionally semi-angular, very shallowly reticulate, shiny, brown; testa not very hard, 70-100 x 50-70 pm, anticlinal walls straight to mostly slightly undulate, inner periclinal wall with numerous small pits. Figure 3.
This new species is characterized by its compact, dense habit, glabrous leaves, white flowers with erect to spreading lobes, pedicel mainly with simple hairs and very few plumose hairs admixed and a capitate-peltate stigma. It is a high altitude species. It is most similar to the widespread and common E. calycina, especially the Cederberg form, which differs in having larger appendages on the anthers (about twice the size), many more plumose hairs on the stem and pedicel, a hairy ovary with truncate stigma, finely hairy leaves, and in growth is woody and erect up to 1 m tall. Several of the collections of the new species were originally identified as being E. calycina.
It is also similar to E. laxa [= E. lucida var. laxa] which differs in having dense plumose hairs on the stem, leaves and pedicel and pink flowers. It is a coastal endem ic on recent sands of the west coast, north of Cape Town. The typical E. lucida is widespread along the mountains of the western region from Cape Town to the Kamiesberg in Namaqualand, and forms sparsely branched, fastigiate shrubs, up to 1.5 m tall.
Erica anemodes is confined to slopes near the sum mits of the highest peaks in the Hex River Mountain complex and adjoining Keeromsberg between Ceres and Worcester, mostly in the range of 1 500-1 900 m in alti tude (Figure 2). Stokoe's collection from 1 200 m is the lowest recorded. The type collection was made near the summit of Waaihoek (= windy comer), hence the epithet (Greek, anemodes = windy). This latter locality was on a rather dry, stony, west-facing slope as opposed to the records of Esterhuysen and McDonald from southern slopes, either rocky and/or peaty or cliffs. Only three of the collections recorded the habit of the species-low compact shrublets with M cDonald noting that the species was locally common. The type collection came from a population which contained numerous small shrublets just in that area, and was made on an excursion with Ms Esterhuysen.
A comment given by Esterhuysen on her collection from the Keeromsberg is significant-'growing with E. calycina' which would indicate that she noted the dis tinctness of this new species in the wild.
It would appear that the species is tending towards wind pollination since the nectaries are very reduced and almost non-existent and the stigma is expanded. Unfortunately the recording of pollen discharge was not noted when Oliver 8793 was collected. ing with continuous apical growth, secondary branches occasional to numerous but not at every node, 10-50 mm long, mostly terminating in an inflorescence, occasional ly with continuous apical growth, tertiary branchlets occasional, intemodes short, 1-2 mm long, sparsely puberulous with simple spreading hairs. Ovary 4-locular, broadly ellipsoid to obovoid, ± 1.2 x 1.0 mm, emarginate, glabrous, nectaries around base; ovules ± 30 per locule, spreading from centrally placed placenta; style ± 1.5 mm long, included; stigma capitate. Fruit a thinwalled dehiscent capsule, broadly obovoid, ± 1.4 x 1.6 mm, valves splitting ± 2/3 their length, and to ± 30°, septa on valves only. Seeds ellipsoid, ± 0.6 x 0.4 mm, slightly reticulate, pale yellow-brown, testa thin with elongate cells ± 100 x 50 (am, anticlinal walls finely undulate, inner periclinal walls densely and finely pitted. Figure 4. Erica viminalis is characterized by its tall growth (3 to 5 ft according to Esterhuysen) with fast-growing wil lowy branches, (hence the epithet, viminalrs = bearing long flexible shoots/branches or willowy), 4-nate leaves with 0-2 sclereids around the midvein, the numerous (± 30) ovules per locule (a large number for a small-flowered species) and the long thin anther appendages.
It is most similar to several species in the section Orophanes. It shares with E. gracilis the 4-nate leaves, the similar corolla and sepal shape and size, and the glabrous ovary and leaves with few sclereids, but that species has much smaller leaves and darker pink flowers and forms a dense, small, delicately branched shrub. With E. leucantha it shares the white flowers, similar anther appendages, glabrous ovary, similar stigma type and 4-nate leaves having few sclereids, but it does not have the glabrous leaves and stems, and the narrow cyathiform corolla. It is also similar to E. sitiens in many of the above characters but that species has much larger flowers with a short pedicel and only 6 ovules per locule and forms smaller fastigiate shrublets up to 1 m tall.
The species is known only from the type collection made by Elsie Esterhuysen in 1959. Her locality is at the base of the "amphitheatre' on the east side of Milner Peak in the Hex River Mountains ( Figure 5). This is the side of the range that faces down into the Hex River Valley and is very dissected with a veritable amphitheatre of very large rocky cliffs and deep gullies. She notes that the plants grew on wet shaded cliffs and that they were conspicuous due to their size. Fortunately, some older flowers with sufficiently matured fruits were also present on the very floriferous branches, which she collected. Although we have not seen the species in the wild, the material reminds us of the rapid growth pattern and habit of two other species we have recently described, namely E. oakesiorum from the Riviersonderend Mountains and E. magnisylvae from near Gansbaai.
The relationships of this new species are rather vague with characters shared with several different groups of species. The nearest allies seem to be the E. leptoclada-E. flacca complex because of the similar prominent thickened midrib especially on the sepals-this vein enlarges considerably in the complex and forms a dis tinctive apical callosity which is missing in E. limnophi la. The hairiness with glands intermixed, the slightly widened filament and the low spreading habit, all agree with this complex, but the differences are the short pedi cel, recaulescent bract and bracteoles approximate to the calyx. In the complex, the bract is characteristically nonrecaulescent (i.e. placed on the main branch) and leaf like and the two bracteoles are placed right at the base of a pedicel which is long, relative to the flower. They also have pollen in monads which is not shared by E. limnophila.
Another group to which the new species is similar is the E. trichoclada-E. cederhergensis group with their often sprawling habit, stalked glands on the leaves and stems, hairy corolla, ovary and appendages and the cap itate to truncate stigma: even the transverse section of the leaves is similar with the very narrow epidermis and numerous sclereids surrounding the midvein-not a very common feature in the genus. The main difference of the new species lies in the fine very short pubescence as opposed to the long villous pubescence in the group.
The glands and hairs on the leaves, corolla and ovary, similar stigma and anther appendages and the similar transverse section of the leaf is shared with the E. pubescens-E. sphaeroidea group but the species in this group form erect stout plants and grow in dry habitats and have multi-whorled inflorescences.
The features of E. limnophila are prostrate, spreading plants, 3-nate leaves, pilose corolla, long, thin anther appendages, short pubescence with long, stout, glandtipped hairs intermixed, inflorescence continuing with vegetative growth often before the flowers have opened and the thickening of the midvein in the leaves, bracts, bracteoles and sepals, but with no apical callosity. The thickening of the vein in the sepals and the singlewhorled inflorescence which continues vegetative growth, are features not seen in any other species of Erica.
Erica limnophila is another new high altitude species, but in this case, is confined to the Dutoitskloof/Wemmershoek Mountains ( Figure 5). It has been recorded from only two localities about 8 km apart. In both local ities its habitat has been recorded as being around the edge of a marsh/swamp/pan, hence the choice of epithet, limn-= pertaining to standing water, pools, lakes, -philus = loving. In this habitat, the species would undoubtedly be a single-stemmed reseeder. Due to the extensive fires in this mountainous region during February/March 1999, fresh material and mature fruits could not be collected. This will have to wait for at least another five years.