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

Three new species of Erica L. from the mountains of Western Cape, South Africa, are described. £. rustieula E.G.H.Oliv. with an indehiscent fruit, is confined to sandy places in the eastern Cold Bokkeveld. E humidicola E.G.H.Oliv. is a highly localized endemic in seepages in the Kogelberg Biosphere Reserve and E rimarum E.G.H.Oliv. is restricted to rock faces at high altitudes in the Hex River Mountains. Erica rustieula E.G.H.Oliv., sp. nov. in grege specierum olim in generibus minoribus fructibus indehiscentibus positorum, Ericae bokkeveldiae E.G.H.Oliv. affinis, sed ab ea antheris quatuor. calcaribus minimis, filamcntis latis, ovario villoso sine nectariis. tubo calycis non crasso, foliis adaxialiter pubescentibus differt. TYPE.—Western Cape, 3219 (Wuppertal): northern Cold Bokkeveld, Kleinveld KM), main valley NE of Sneeukop. 1 240 m. (-CD). 6 May 2(XK). Oliver 11517 (NBG. holo.; BM. K. MO. NY. PRE. S). Shrub compact rounded, 100—150( —300) x 100-300 mm, erect to semi-spreading, single-stemmed reseeder. Branches: numerous main branches of short growth 10-15 mm long, terminating in an inflorescence or con­ tinuing vegetative growth, each with 2-5 short, recurved secondary branchlets, 2-10 mm long, not from each node, intemodes very short. ± 1 mm or less, all covered with dense, very short, white reflexed hairs. Leaves 3nate. imbricate. ± 1.6-2.0 x 1.0 mm. adpressed. elliptic, subacute, adaxially shortly hairy and flattened, abaxiallv glabrous and rounded, margins shortly ciliate when young, sulcus narrow and closed at base: petiole adpressed. ± 0.4 mm long, glabrous, shortly ciliate. Inflorescence: flowers 3(6)-nate in 1(2) w horls at ends of main and secondary branches, nodding, umbel-like w hen 2-whorled: pedicel 0.3 mm long, glabrous, pinkish red: bract partially recauleseent and approximate to calyx. ± 1.0 x 0.5 mm. elliptic to oblong, subacute, pale pink to pink and sometimes green-tipped, glabrous, margins shortly ciliate and with a few subsessile. dark red nonsticky glands towards apex, sulcus small, narrow: bracteoles 2 approximate and adpressed to calyx, otherwise like the bract. Calyx 4-lobed: ± 1.5 x 2.2 mm. shortly and broadly funnel-shaped, hard and wax-like, pink: tube ± 1 mm long; lobes adpressed to corolla, ± 0.8 x 1.0 mm. broadly elliptic, subacute, sulcus 0.2-0.3 mm long, nar­ row. Corolla 4-lobed, 2.5 x 2.5-3.5 mm. shortly and widely funnel-shaped, glabrous, pink, with thin semi­ transparent broad tube I mm long; lobes semi-spreading. ± 1.5 x 2.0 mm. broadly ovate, rounded or subacute. * Compton Herbarium. National Botanical Institute. Private Bag X7, 7735 Claremont, Cape Town. MS received: 2000-07-03. margins entire, with sessile, dry, dark-red glands and short reflexed hairs. Stamens 4. subexserted, free: fila­ ment broad ± 2.5 x 0.5 mm. oblong, straight to slightly curved apically. glabrous, w hite: anthers bilobed. ± 0.8 x 0.7 mm. obovate in outline in adaxial view, dorsally attached near base, minutely appendiculate. spurs very small. ± 0.1-0.3 mm long: thecae ± 0.8 x 0.35 mm. ovate in lateral view, erect, with a few papillae, otherwise glabrous, almost black: pollen in monads. Ovary 2-locular. ± 0.7 x 0.6 mm, ovoid, complanate. obtuse, upper three quarters covered with dense straight long hairs, lower quarter glabrous, septum very thin and fragile, nectaries absent: ovules 1 per locule. pendulous from a subapical placenta: style exserted. ± 5 mm long, filiform, curved, glabrous, reddish pink: stigma simple-truncate to slightly widened. Fruit indehiscent. ± 1.2 x 2.0 mm. ovoid, pericarp very thin and papery with exocarp thin, mesocarp composed only of numerous crystals and endocarp thin and brow n. Seed one per fruit. ± 1.2 x 1.8 mm. ovoid, testa thin, very shallowly reticulate, vellow brown; cells 25-50 x 75-100 |im, irregularly elongate with slightly thickened jigsaw-like anticlinal walls, numerous small pits in inner periclinal w alls. Figure 1. This new species was discovered just as the revision of the 84 species of indehiscent-fruited Erica was in press (Oliver 2(XX)). The species is at first sight very similar to Erica hokkeveldia E.G.H.Oliv. in the structure of the shrublet and of the flower—the pink, open flowers w ith dark exserted stamens and very long style, the petaloid pink bract, bracteoles and calyx and the 2-locular ovary w ith a single ovule in each locule. A close examination of the flowers reveals that it differs from that species in having only four stamens (not eight), minute spurs on the anthers (not long, broad and serrated spurs), oblong, broad filaments (not linear and thin), an ovary covered all over with long thin hairs (not shortly hairy at the apex only) and lacking any nectaries, and the corolla tube thin, delicate and semitransparent (not hard and thickened). E hokkeveldia w as included in the former genus Eremia as Eremia calycina Compton (Oliver 1976). There are also slight differences between the two species in other organs—the leaves in E. rustieula are 148 Bothalia 30,2 (2000)

The species is at first sight very similar to Erica hokkeveldia E.G.H.Oliv. in the structure of the shrublet and of the flower-the pink, open flowers w ith dark exserted stamens and very long style, the petaloid pink bract, bracteoles and calyx and the 2-locular ovary w ith a single ovule in each locule.A close examination of the flowers reveals that it differs from that species in having only four stamens (not eight), minute spurs on the anthers (not long, broad and serrated spurs), oblong, broad filaments (not linear and thin), an ovary covered all over with long thin hairs (not shortly hairy at the apex only) and lacking any nectaries, and the corolla tube thin, delicate and semitransparent (not hard and thickened).E hokkeveldia w as included in the former genus Eremia as Eremia calycina Compton (Oliver 1976).
There are also slight differences between the two species in other organs-the leaves in E. rustieula are Bothalia 30,2 (2000) 4mm N , lOOpm pubescent adaxially (not glabrous), narrower sepals which are broadly elliptic and subacute (not very broad ly orbicular and broadly obtuse).In the seeds the testa is slightly thicker, being yellowish with slightly thickened, more jigsawed anticlinal walls and numerous small pits in the periclinal walls (not transparent with undulate non thickened anticlinal walls and large-pitted periclinal walls in E. bokkeveldia).
The texture of the corolla tube is significantly differ ent in the two species.In E. bokkeveldia the hard tube envelops the delicate indehiscent fruit and remains around it as a protection when the whole flower is shed onto the ground.The fruit is extremely delicate with a very thin, parchment-like pericarp and almost non-existent, transparent testa.In E. rusticula this tube is not hard and therefore does not provide the same protection to the fruit which is, however, a bit tougher with a slightly thicker pericarp and with a slightly thickened seed testa.
Erica bokkeveldia is a somewhat isolated species in the genus and is hypothesized to be allied to E. cetrata E.G.H.Oliv.(Oliver 2000).We would suggest that E. rus ticula also belongs in this alliance.The position of these species within the genus Erica, which has now been con siderably enlarged with the inclusion of all the indehiscent-fruited genera (Oliver 2000), is unresolved due to the lack of any clear indications of a 'natural' infra generic division of the genus.All the indehiscent-fruited species are retained at the end of the genus as an interim measure.E. rusticula is therefore placed with E. bokkeveldia as species number 45a (Oliver 2000).
The above three species are known from the Ceres District with E. cetrata being more widespread to the base of the Hex River Mountains and eastwards to the Bonteberg near Touws River.The other two are Cold Bokkeveld endemics.E. rusticula is confined to the rocky ridges between Bokkeveld Sneeukop and Bloukop in the northeastern areas, whereas E. bokkeveldia occurs at lower altitudes just further south and to the west-they do not co-occur.Both inhabit sandy or sandy-gravelly flat areas where the other shrubs are also very low.
Erica rusticula has been found in five populations on the Farm Kleinveld 100, hence the epithet-rusticulus = of the countryside [diminutive form], 'Klein-veld'.This farm covers the areas east of the Skurweberg Range from Bokkeveld Sneeukop to Bloukop (Figure 2).In the higher altitude populations the associated vegetation is currently mature, rather old and undisturbed, whereas at the type locality there is a fair amount of wind erosion and farming disturbance on the pure quartzitic sandy flats.In the latter area the plants are extremely old with very gnarled woody stems.Unfortunately, due to the extremely dry summer of 1999/2000 most of the popula-tions were not flowering well except for the population at the northeastern base of Sneeukop, hence its selection as the type.The species was noted among Erica materi al collected for our interest by a team from the Protea Atlas Project for which we are always very grateful.
Apart from variation in the size of the plants and their flowering condition between the populations, no varia tion in morphological characters was noted.
The lack of nectaries in E. rusticula is an unusual fea ture since there is no accompanying enlargement of the stigma that is usually associated with this condition.The nectaries are clearly present in E. bokkeveldia and bees have been seen visiting a population of the species in Hartebeeskloof.No pollinating insects were noted visit ing the plants of E. rusticula and there was no indication of pollen being shed when the plants were manually dis turbed.The pollination syndrome needs to be investigat ed in the light of the differences between the two species.Flowering time: April and May.This is a very dry period of the year in the area.However, in good years the nor mal early autumnal rains around the Easter period could be very beneficial and trigger the flowering of the species.This flowering time is in contrast to the September/October flowering period for E. bokkeveldia.Shrub erect, 0.5-1.0m tall, bushy in open areas or long and lanky in thicker, older vegetation, single stemmed reseeder.Branches: a few main erect ones, 100-300 mm long, mostly with continuous apical growth, numerous secondary branches, 10-20 mm long at every node, few, very short, 1 ^ mm long tertiary branchlets, intemodes on main branches 8-10 mm long, all branches with short spreading hairs.Leaves 4-nate, ± 3.5 x 0.6 mm subspreading, adaxially flattened, abaxially rounded, hirsute all over, sulcus narrow and closed at base; petiole 0.6 mm long, adpressed, glabrous, ciliate.Inflorescence: flowers 1-3 at ends of short lateral sec ondary and tertiary branchlets crowded along main branches into loose spike-like synflorescence; pedicel 2 mm long, pubescent, red; bract partially recaulescent about quarter to third way up pedicel, ± 0.6 x 0.3 mm, ovate-triangular, subacute, puberulous, ciliate, with small subapical sulcus, pink; bracteoles 2, positioned slightly higher than bract, slightly longer than bract, oth erwise similar.Calyx 4-partite, segments ± 1.5 x 1.2 mm, broadly ovate-elliptic, acute, adpressed to corolla, short ly and narrowly sulcate up to quarter their length, finely puberulous, shortly ciliate, reddish pink.Corolla 4lobed, ± 4.5 x 3.0 mm.broadly campanulate, very finely puberulous, dark pink: lobes ± 2.0 x 1.5 mm, broadly rounded, entire, subspreading to spreading, devoid of hairs towards margins abaxially.Stamens 8, included or just manifest, free; filaments ± 2.5 mm long, linearoblong, straight or with slight S-bend, glabrous, white; anthers bifid, dorsally attached near base, erect, sub quadrate in adaxial view, appendiculate, spurs ± 0.5 mm long, pendulous, slightly decurrent along apex of fila ment, thick, entire, sparsely papillate, golden; thecae dark brown, slightly parted but erect, ± 0.8 x 0.4 mm, oblong-elliptic with slight nose in lateral outline, obtuse, papillate-strigose on adaxial margins; pore about half length of theca; pollen in tetrads.Ovary 4-locular, ± 1.3 x 1.1 mm, broadly obovoid.emarginate, with small basal nectaries, hirsute; ovules 4 or 5 per locule, spreading from placenta centrally placed on axis; style exserted, 3 mm long, cylindrical, glabrous; stigma capitellate.Fruit a dehiscent capsule, ± 4.5 x 4.0 mm, valves free almost to base but not spreading much, septa about 80% on valves and 20% on columella.Seeds ± 0.5 x 0.4 mm, subspheroid-ellipsoid, shallowly reticulate, golden brown; cells subequally subquadrate, ± 45 x 55 (im, with slight ly undulate anticlinal walls and small pores in inner periclinal walls.Figure 3.This new species falls within a large group of waterloving species with 4-nate leaves such as the large tubular-flowered Erica macowanii Cufino and the common small-flowered E. parviflora L. They all have in com mon, leaves with a thin cuticle, large epidermis and few sclereids (< 5), nearly all nodes on the main branches bearing secondary branchlets, terminating in 1-4-flowered inflorescences, the main branches continuing with vegetative growth; corolla usually hairy and often finely so as in E. feminarum E.G.H.Oliv., E.fervida and E. pillansii.Most species in the alliance have no anther appendages-the few exceptions being E. parviflora L., E. velitaris Salisb., the E. fervida/pillansii complex and this new species, and most with no hairs on the ovary.

Paratype material
The closest relatives of E. humidicola appear to be the showy scarlet-flowered E. fervida and E. pillansii, which form a complex and are both also confined to the Kogelberg Biosphere Reserve.It can, however, be distin guished by the pink colour of its flowers which have a shorter open campanulate corolla (not long, tubular-campanulate to tubular) and appendiculate anthers which have papillate hairs on them (not smooth muticous anthers).The species is placed here in §Evanthe with these two species despite the very short corolla tube, the others being in §Ephebus (E.parviflora) and §Orophanes (E. velitaris).This clearly points to the unsatisfactory subgeneric system mentioned above under E. rustieula.
The species is very restricted in its distribution being known thus far from only two small marshy/seepage areas in Spinnekopsneskloof below Dwarsrivierberg (Figure 2).In both populations the species grows in stands of much taller, erect shrubs of species of Leucadendron (Proteaceae) and Psoralea (Fabaceae).In the northern population it occurs as the only heath, where as the southern population is dominated by taller plants of the yellow-flowered E. campanularis Andr.In both there are few plants, perhaps only several dozen in each.Two other species which inhabit damp/wet places were noted nearby, the small pink-flowered E. intervallaris at the edges of the seeps and the tall, orange tubular-flowered E. curviflora alongside the stream near the northern popula tion.Flowering time: September-October.
A single plant of undoubted hybrid origin between E. humidicola and E. campanularis was recorded in the southern population by Mrs Amida Johns of the Kogelberg Biosphere Reserve who accompanied us on the investigation of the new species.The hybrid had paler pink flowers with whitish tips and the narrower campanulate corolla shape of E. campanularis.
The name is derived from the habitat preferencehumidus = damp, wet, incola or -cola = dweller.
The species was brought as a small branchlet by Ion Williams in 1964 and again in 1969 by former colleague, Charlie Boucher, during his survey of the Kogelberg Reserve.It is the second species recorded as endemic to the Spinnekopsneskloof, the other being E. vallis-aranearum E.G.H.Oliv.which occurs at 1 000 m at the head of the kloof on steep south-facing slopes and which is extremely rare.Shrub compact, gnarled and twiggy, 40-100 mm tall, single-stemmed, stem very old and woody.Branches: many main branches ± 10 mm long, erect or recurved, mostly terminating in an inflorescence, occasional sec ondary branchlets up to 5 mm long, all glabrous.Leaves 2-nate, imbricate, ± 3.0 x 0.7 mm, erect, lanceolateoblong, adaxially concave, abaxially rounded, with acute margins, glabrous, with a few non-sticky short-stalked glands on basal margins and petiole, apex yellow-cuspi date, sulcus very narrow and closed at base; petiole adpressed, ± 0.5 mm long, glabrous.Inflorescence: flow ers 2-nate in one whorl at ends of main branches; pedicel ± 2.0 mm long, curved at base, glabrous, reddish; bract partially recaulescent, about one third up pedicel, ± 2.0 x 0.6 mm, lanceolate, acute, glabrous, maroon, margin in lower half with sessile or very short-stalked, non-sticky, dark red glands, sulcus narrow, about one third its length; bracteoles 2, attached about two thirds up pedicel, simi lar to bract.Calyx 4-partite, maroon; segments adpressed to corolla, imbricate, ± 2.5 x 1.1 mm, ovate, acute, glabrous, margins with small, sessile or short-stalked, non-sticky, dark red glands, mainly in lower half, sulcus narrow, about one third length of segment.Corolla 4lobed, ± 3.0 x 2.5 mm, cyathiform, maroon, thick in tex ture, glabrous; lobes 1.5 x 1.0 mm, rounded, entire, erect.Stamens 8, included, free; filament ± 1.1 x 0.2 mm, broadened towards base, with small apical sigmoid bend below anther, glabrous, white; anthers ovate in adaxial view, bilobed, dorsally attached, appendiculate, spurs pendulous, ± 0.5 x 0.1 mm [as long as theca], serrate and finely ciliate; thecae ± 0.5 x 0.4 mm, broadly ellipticrhombic in lateral view, dark brown, glabrous; pollen in tetrads.Ovary 4-locular, ± 0.9 x 1.0 mm, broadly ovoid, emarginate, with well-developed nectaries around base, glabrous; ovules ± 15 per locule, spreading from placen ta, central on axis; style included, 1.1.mm long, broadly cylindical, tapering towards base; stigma capitate.Fruit a dehiscent capsule, valves splitting about 70% with septa on valves only and free from columella.Seed ± 0.5 x 0.3 mm, unequally ellipsoid with flat and rounded sides, light brownish orange, shallowly reticulate; testa thin, cells elongate, ± 100 x 30 [im, irregularly elliptic, with slightly raised, jigsawed anticlinal walls and dense ly pitted, inner periclinal wall.Figure 4.This new species is most similar to E. brevicaulis Guthrie & Bolus and the recently described E. navigatoris E.G.H.Oliv.(Oliver & Oliver 1998) both of which occur in the same mountain range.In growth and habitat and flower colour it is most similar to the former which has hairy margins to the leaf, a hairy ovary and short anther appendages, which are lateral on the filament.
Despite the erect, much larger size of the plants and flowers of E. navigatoris, the new species is probably more closely related to it-it is like a miniature version of that species.They share glabrous branches, cuspidate leaves, non-sticky, slightly stalked red glands on the edges of the bract, bracteoles and sepals which are all similar in shape and position and the anthers with broad irregularly serrate and ciliate appendages.E. rimarum differs, however, in a number of characters-2-nate leaves (not 3-nate), corolla cyathiform and maroon (not urceolate and white), anthers included (not manifest), stigma included (not exserted) and anther appendages on the anther theca (not along the apex of the filament).
Erica rimarum forms small, gnarled, woody shrublets, growing on south-facing ledges and clefts on large rocks or cliff-faces, hence the epithet, rima = a cleft or fissure.The species is widespread on the peaks in the southwestern part of the Hex River Mountain range from Milner Peak to Waaihoek, where it grows at high alti tudes (Figure 5).Miss Esterhuysen showed the first author the plants growing on the northern slopes west of Milner Peak and at the time we discussed its probable identity as E. brevicaulis.Many collections have been made by her and also by Thomas Stokoe, both intrepid collectors of high-altitude species, often growing in inac cessible rocky places in these rugged mountains.The last collection of the species was that by Esterhuysen and the first author in 1960.The record from Stettynsberg is interesting, since no collections have been made in any of the intervening mountains of the Du Toitskloof com plex.Flowering time: October-December.
The species possesses large nectaries around the base of the ovary, which feature should indicate that the small flowers are pollinated by insects.
Johns 38 (reference collection at Kogelberg Reserve and at Harold Porter Botanical Garden, Betty's Bay).