Distribution

Indonesia: West and South Kalimantan.

Ecology

Lakeside rain forest Flowering: October.

The type specimen of Vaccinium adenopodum was not seen by Vander Kloet (Reference Vander Kloet2005), who excluded it from his treatment. Examination of the type specimen shows that the basal glands are clearly (to c.1 mm) although not strongly prominent. Sleumer had described them as “generally not properly prominent” (Sleumer, Reference Sleumer and Van Steenis1966–1967). The disc is densely hairy, not glabrous as Sleumer described it. I could not see any corollas, so my description of the flowers follows that of Sleumer (Reference Sleumer and Van Steenis1966–1967). I have moved Vaccinium megaphyllum Sleumer var. adenophorum Sleumer into synonymy with Rigiolepis adenopoda. Vaccinium megaphyllum is a poorly known taxon, as the flowers have not been described. The vegetative description fits well, and the locality is close. Giesen 65 (L) agrees reasonably well except that the bracts are minute (c.1 mm) and much shorter than the pedicels. Another Leiden specimen without locality, accession number 951 304 380, agrees well vegetatively but has much longer, laxer inflorescences.

This species perhaps comes closest of the Bornean material to the widespread Vaccinium acuminatissimum Miq., with its protruberant marginal leaf glands. It has larger, shinier leaves than is usual in Vaccinium acuminatissimum, however, and only laxly distributed simple glands on the undersides of the leaves, unlike the usually dense, complex glands of V. acuminatissimum.

Fig. 5.

Rigiolepis andersonii (Sleumer) Argent. RBGE accession number 19781702; from G. Api, Sarawak, the type locality. A, Flower (scale bar, 2 mm), with inset showing surface detail (scale bar, 0.5 mm); B, stamens (scale bar, 1 mm); C, style with disc (scale bar, 1 mm); D, habit (scale bar, 3 cm). (Illustration: Claire Banks.)

Fig. 6.

Rigiolepis andersonii (Sleumer) Argent. Flowering plant, Gunung Api, Sarawak. (Photograph: G. Argent.)

Epiphytic shrub to 1 m. Twigs minutely puberulous; lateral buds 2 × 1 mm, mostly less than half the length of the petiole, arising distinctly above the leaf axil, the scales covered with curly hairs. Leaves: petiole 4–5 × 4–5 mm, shallowly grooved, minutely puberulous; blade 50–80 × 30–45 mm, ovate to ovate-elliptic, base truncate, rounded, to subcordate, margin entire, flat, basal glands impressed next to petiole, smooth and glossy above, glabrous except for minute glandular hairs beneath, with 6–8 arching lateral veins, a basal pair and progressively more pinnate ones distally, slightly prominent above, more distinctly so beneath when dry, midvein depressed for its whole length above in dry specimens, prominent beneath, reticulation densely prominent, especially beneath, apex obtusely acuminate for 5–10 mm. Inflorescence: the racemes laxly many-flowered; rachis 50–80 mm, slender, puberulous and with brown glands; bracts small c.1 mm long, acicular; bracteoles to c.1 mm long, glandular hairy, one basal, one about halfway up the pedicel. Flowers: pedicels 5–6 mm, slender, puberulous and glandular hairy; calyx 1–1.5 × c.2 mm, cup-shaped, tube subtruncate at base, covered in brown glandular hairs, lobes acutely triangular, glandular, fimbriate and ciliolate; corolla c.4 × 4.5 mm, urceolate, pale yellow, strongly scented, pubescent and glandular muriculate especially at angles outside, nearly glabrous inside, lobes c.1 mm, reflexed; stamens: filaments 1.5 mm, linear, densely hairy; anther cells c.0.5 mm, ovoid, echinulate; tubules 1 mm, narrow; disc subdensely short-hairy; style slender, laxly short-hairy in proximal 3/4 or nearly glabrous. Fruit globose, c.5 mm in diameter.

Distribution

Malaysia: Sarawak. Known only from the environs of Mount Api, but this species is likely to occur on Mount Benarat and other associated limestone peaks in the Mount Mulu National Park.

Ecology

In submontane forest, epiphytic and on peat overlying limestone 915– 1220 m. Flowering: July, probably at other times as the plants in cultivation flower several times a year.

Conservation assessment

LC. This species is common on Mount Api in the Gunung Mulu National Park, which is a protected area. Although the summit area is prone to droughts (and fire, to judge by the name of the mountain), the deep grikes afford a variety of habitats and protection from the worst effects of desiccation.

Very similar to Rigiolepis salicifolia but with shorter broader leaves. Vander Kloet (Reference Vander Kloet2005) says [leaf] “blade glabrous underneath”, while Sleumer (Reference Sleumer and Van Steenis1966–1967) says “glabrous except for fine, hardly visible setular hairs beneath”. In fact, the leaves are minutely glandular hairy when very young, leaving only a very indistinct punctate surface. Cultivated from plants growing in the type locality (Mount Api) at the Royal Botanic Garden Edinburgh, where it maintains its distinctively shaped leaves. The original description is modified from observations of the cultivated specimens.

3. Rigiolepis balgooyi Argent, sp. nov.

Rigiolepis balgooyi is similar to Rigiolepis winkleri in having long glandular hairs which overtop the simple hairs on the inflorescence but differs in its lowland habitat, protruding marginal leaf glands, shorter acuminate leaf apices and larger stamens with filaments much longer than the anthers. – Type: Indonesia, West Kalimantan, c.100 km south of Pontianak, Gunung Palung Nature Reserve, 14 vi 1986, Balgooy & Setten 5433 (holo BO, iso L). Figs 4, 7.

Fig. 7.

Rigiolepis balgooyi Argent. Isotype, Balgooy & Setten 5433, Gunung Palung, West Kalimantan. (Scan: Robyn Drinkwater.)

Epiphytic shrub. Twigs very shortly patently hairy and with some short brown glandular hairs, leaves laxly complanate; lateral buds c.5 × 3 mm, broadly subulate, about as long as petiole, densely hairy. Leaves: petiole 5–6 × c.1 mm rugose, puberulous; blade 55–85 × 20–35 mm, sulcate, narrowly ovate to elliptic, base rounded, margin slightly recurved, with a distinctly protruding gland on each side c.1.5 mm from petiole, lamina glabrous above, with simple glandular hairs beneath, with 2 lateral veins on each side, the first from the base disappearing about mid-leaf, the second emerging a few millimetres above base, arching and ascending to well above mid-leaf, main veins impressed above, prominent beneath, reticulation obscure, apex shortly acuminate (1–1.5 mm). Inflorescences mostly of solitary racemes, sometimes paired; rachis slender, 10–15 mm, with several triangular to subulate stipule-like bracts at base, moderately densely hairy with curled simple hairs and scattered long-stalked glandular hairs overtopping simple hairs; bracts c.1 mm, ovate-acuminate, with both simple and glandular hairs; bracteoles minute, subulate, with both simple and glandular hairs. Flowers: pedicels slender, 6–9 mm with simple and glandular hairs similar to those on rachis; calyx tube very short, 0.5 × 0.7 mm, densely shortly pubescent with both simple and glandular hairs, lobes c.0.6 mm, narrow-triangular; corolla c.4 × 2.5 mm, urceolate, pale yellow, with a few short simple and glandular hairs, mostly at angles, glabrous inside, lobes c.0.3 mm, reflexed; stamens: filaments 2.2 mm, wavy, linear, hairy; anthers cells 1.5 mm, oblongoid, papillose, 0.8 mm; tubules slender, erect, introrsely slit almost their entire length, 0.9 mm; disc laxly hairy on top; style 3.2 mm, slender, glabrous. Fruit not seen.

Distribution

Indonesia: West Kalimantan. Known only from the type specimen.

Ecology

Epiphytic in lowland dipterocarp forest on alluvial soil at c.30 m elevation. Flowering: June.

Etymology

Named after Max Michael Josephus van Balgooy, for his outstanding contribution to Southeast Asian botany.

Conservation assessment

LC. This species occurs in Gunung Palung Nature Reserve, which should afford it some protection.

Similar to Rigiolepis leptantha var. elliptica (f. elliptica of Sleumer, Reference Sleumer and Van Steenis1966–1967) in its lowland habitat and general leaf shape, but that taxon has smoother leaves, only a single pair of faint lateral veins, and impressed not protruding marginal leaf glands, and it does not have the long-stalked glandular hairs on the inflorescence.

4. Rigiolepis bigibba (J.J.Sm.) J.J.Sm., Blumea 1: 336 (1935). – Vaccinium bigibbum J.J.Sm., Bull. Jard. Bot. Buitenzorg, ser. 3, 1: 408, t. 55 (1920). – Type: Indonesia, West Kalimantan, Lanjak, P. Pandjang, 18 vii 1874, J.E.Teysmann 7966 (lecto BO, designated by Sleumer, Reference Sleumer1961; isolecto FI, L). Figs 4, 8.

Fig. 8.

Vaccinium bigibbum (≡ Rigiolepis bigibba). Type plate: Tab. 55., reproduced from the Bulletin du Jardin Botanique de Buitenzorg, series 3e, volume I.

Shrub to 5 m. Twigs puberulous, the leaves laxly complanately arranged; lateral buds 3–4 × c.2 mm, broad, scales about half as long as petiole, densely hairy. Leaves: petiole 6–8 × 2–3 mm rugose, puberulous; blade 75–21 × 30–80 mm, ovate to ovate-elliptic, base obtuse to rounded, margin slightly recurved; contracted and decurrent in distal 1/3 of petiole for c.2 mm where there is a thick protruding gland on each side, lamina glabrous except for some puberulence at base and with dense minute early caducous glands beneath, with 4–8 strong high-arching lateral veins from and from somewhat above base (outer pair of veins less distinct), main veins impressed although raised within a shallow depression above, sharply prominent beneath, other shorter veins from upper part of midvein in several pairs, irregular and much less distinct on both sides, reticulation lax, sometimes slightly impressed, but mostly prominent on both sides, occasionally less visible beneath, apex longer (to 30 mm) or more shortly caudate-acuminate, subacute. Inflorescence of 2 or 3 racemes in axillary fascicles, or solitary; rachis 20–35 mm, robust, with several triangular to subulate stipule-like bracts at the base, laxly or densely shortly pubescent and finely glandular hairy; bracts c.2 × 1 mm, ovate-acuminate, with simple and glandular hairs, more than half the length of pedicels; bracteoles minute, subulate, with both simple and glandular hairs. Flowers: pedicels thick, 2–3 × 1.5–2 mm; calyx tube very short, c.0.7 mm long, densely shortly pubescent and finely glandular hairy, lobes c.0.6 mm, narrow-triangular, tips red; corolla c.4 × 2.5 mm, urceolate, cream or yellow, 5-angular, slightly puberulous with simple hairs outside, glabrous inside, lobes c.0.3 mm, reflexed; stamens: filaments 2 mm, wavy, linear, hairy; anther cells c.1 × 1 mm, papillose; tubules c.1 mm, slender, erect, introrsely slit almost their entire length; disc densely hairy; style 4 mm, slender, glabrous. Fruit not seen.

Distribution

Brunei. Indonesia: West and Central Kalimantan. Malaysia: Sarawak.

Ecology

On boulders in a forested ravine, epiphytic in peat-swamp and Dipterocarpus forest, mostly at low elevation but up to 1700 m. Flowering: May to August.

This species is somewhat problematic taxonomically, and I have restricted it to forms with relatively short rachises and pedicels (as in the original description of Smith, Reference Smith1920) and with relatively long subtending bracts which are over half as long and often as long as the rather robust pedicels (not slender as Sleumer, Reference Sleumer and Van Steenis1966–1967, describes them). This better distinguishes Rigiolepis bigibba from R. sulcata rather than the relative hairiness of the leaves, as did Sleumer (Reference Sleumer and Van Steenis1966–1967). Rigiolepis sulcata possesses longer, slender pedicels with the subtending bracts much less than half the length of the pedicels, as does R. irawatiae, which has even smaller bracts. Rigiolepis irawatiae also has glandular hairs overtopping any simple hairs on the inflorescence and much smaller basal leaf glands which do not form auricles as they do in the present species and R. sulcata. Nevertheless, Rigiolepis bigibba, R. sulcata and R. irawatiae form a group of species with complex variation of leaf shape, degree to which they are sulcate or smooth, and the multicellular hairs which usually can be seen only with a lens as a yellowish, mealy indumentum.

Smith (Reference Smith1935), in a comment on the illustration of the type (see Fig. 8), says: “It seems that the flower described. . . was somewhat disturbed. In other flowers I found the anther tubules tapering and the spurs well-developed and subulate”. This illustration does, however, nicely show the relative lengths of bract and pedicel.

5. Rigiolepis borneensis Hook.f., Icon. Pl. 54, t. 1160 (1876). – Type: Malaysia, Sarawak, Lobb s.n. (holo K). Figs 3, 9, 10.

Fig. 9.

Rigiolepis borneensis. RBGE accession number 19910076. A, Flower (scale bar, 2 mm); B, stamens and style with disc (scale bar, 1 mm); C, bare branch with inflorescences (scale bar, 1 cm); D, habit (scale bar, 5 cm). (Illustration: Claire Banks.)

Fig. 10.

Rigiolepis borneensis. RBGE accession number 19943021; in cultivation at RBGE, collected from Gunung Serapi, Sarawak. (Photograph: G. Argent.)

Vaccinium borneense W.W.Sm., Notes Roy. Bot. Gard. Edinburgh 8: 329 (1915). – Type: Malaysia, Sarawak, 4 xi 1913, Native collector 66 (lecto E E00412436; isolecto E E00412437, K, SAR).

Climbing or twining shrub to 4 m, with swollen woody base. Twigs thick, covered with long white hairs to 0.4 mm; lateral buds to 3.5 mm, narrowly pointed, mostly as long as petiole, excluding decurrent leaf base. Leaves: petiole, 3–5 × 1–1.5 mm, pubescent with simple white hairs; blade 100–160 × 20–50 mm, elliptic to subovate-elliptic, base acute to obtuse, with decurrent flanges, margin entire, weakly revolute, impressed marginal glands 5–8 mm from petiole, apex acuminate to subcaudate-acuminate, acute, acumen 10–20 mm long, distinct, glabrescent on both sides except for major veins which have long white hairs particularly adaxially, midvein and inner pair of veins deeply impressed above and prominent beneath, with 2 or 4 high-arching lateral veins, other transverse veins numerous, anastomosing, forming a dense reticulation on both sides and often a characteristic scalariform pattern. Inflorescence of racemes from upper axils, many-flowered; rachis 10–25 mm, thick, laxly glandular hairy bracts ovate-acuminate, prominently many ridged lengthwise (costulate) at back, glandular hairy, 2.5–3 × 1.5–2 mm; bracteoles inserted in middle of pedicel, elliptic, prominently ribbed, 1.5–2 × 1 mm. Flowers: pedicels slender, 2–3 mm long, laxly glandular hairy; calyx glandular hairy, deeply 5-lobed, tube densely long hairy, with simple hairs, becoming subglobose, lobes 1.7–2.5 × c.1 mm, ovate-triangular to oblong-ovate, very prominently parallel ribbed when dry; corolla 2.5–3.5 × 1.5–2 mm, subovoid-urceolate, contracted at both ends, white sometimes greenish-white or cream, 5-angled, laxly pubescent outside, lobes 0.5 mm, spreading; stamens: filaments linear, papillose, 0.7–1.2 mm; anther cells ovate-oblong, echinulate, 0.4–0.6 mm; tubules 0.4–0.6 mm, cylindrical, erect; disc pubescent; style glabrous, 2–3 mm. Fruit not seen.

Chromosome number

2n = 24 (Atkinson et al., Reference Atkinson, Jong and Argent1995).

Distribution

Indonesia: West Kalimantan. Malaysia: Sarawak.

Ecology

In low wet forest, or kerangas (heath forest); also recorded from a ridge top, 300–1065 m. Flowering: October to January, mostly from February to March in cultivation, probably irregular in the wild.

Conservation assessment

The low-elevational distribution of this species makes habitat destruction a distinct threat.

See discussion under Rigiolepis above. This is the oldest name in the group of species characterised by distinctively costate bracts and bracteoles, best seen when dry, bracts as long as or longer than the pedicels, and relatively long calyx lobes. The type specimen has a pencil note appended to it, stating “flowers red”. All the associated material has white to pale green flowers which sometimes turn reddish in old age. Truly red flowers in this species would seem doubtful.

6. Rigiolepis chaii Argent, sp. nov.

Most similar to Rigiolepis lobbii var. lanceifolia but differing by calyx lobes c.1 mm long (versus > 2 mm long), calyx lobes and bracts smooth (not costate when dry), and stem indumentum of minute, strongly appressed, curled hairs to 0.1 mm long (not the longer, straight, irregularly patent hairs to 0.4 mm long of R. lobbii var. lanceifolia). – Type: Malaysia, Sarawak, Miri Division, Gunung Mulu N.P., Mentawai, 7 x 1977, Chai S.39726 (holo SAR, iso K). Figs 4, 11.

Fig. 11.

Rigiolepis chaii Argent. Holotype, Mulu N.P., Chai S.39726. (Scan: Robyn Drinkwater.)

Pendulous epiphyte. Twigs densely covered with short, curled, matted hairs to 0.1 mm long; lateral buds to 5 mm long, scales finely acicular, much longer than petioles, patently long-hairy. Leaves: petiole 1–1.5 × 0.5 mm, pubescent with curled hairs similar to those on stems; blade 35–80 × 5–10 mm, linear to linear-lanceolate; base rounded, margin entire, strongly inrolled, basal marginal glands distinct and impressed, c.2 mm from petiole, extreme apex rounded, lamina convex, with scattered brown, clavate, glandular hairs beneath, midvein impressed above, strongly raised beneath, and with numerous, inconspicuous lateral veins, but lacking prominent arching lateral veins, upper surface finely and distinctly reticulate. Inflorescence of solitary, axillary flowers. Flowers: pedicels to 2 mm long from a few small, hairy acicular basal bracts; calyx lobes c.1 × 1.5 mm, with a distinct gland subapically outside; corolla not seen; disc densely hairy with short semi-appressed hairs. Fruit (immature) subspherical, scabrid, c.4 × 4 mm.

Distribution

Malaysia: Sarawak, Miri Division, Mount Mulu National Park. Known only from the type collection.

Ecology

Epiphyte hanging over the river from Sandoricum koetjape (Burm.f.) Merr., in the Setap Shale area. The elevation is not recorded but will be low, probably c.300 m. Flowering time not recorded.

Etymology

Named after Paul Piang Kong Chai, collector of this plant and for his contribution to Southeast Asian botany, particularly for his support for the Royal Geographical–Forest Department Kuching expedition to Mount Mulu, 1977–1978.

Conservation assessment

LC. This species has the protection of occurring in a designated national park.

This species is very distinct despite not being collected in flower, with its long very narrow linear leaves, which are broader than those of the other linear-leaved species and possessing subsessile flowers lacking rachises. The specimen at K has floral remnants glued to it which do not belong to this species!

The new species is superficially similar to some specimens of Rigiolepis lobbii but lacking the straight hairs, costate bracts and calyx lobes (when dry) of that species.

7. Rigiolepis irawatiae Argent, sp. nov.

Most similar to Rigiolepis sulcata but with the dendroid hairs on the leaves being filamentous and scabrous (not lamellar) and the leaves almost smooth, with only a slightly protruding pair of basal glands (to c.1 mm), these at the precise junction between the non-decurrent lamina and petiole; petiole and leaves hairy with multicellular and simple hairs. The rachis is slender, elongate, 60–110 mm (versus up to 50 mm long in Rigiolepis sulcata), and the corollas are densely and completely glandular hairy outside without the simple hairs found in R. sulcata. – Type: Indonesia, Central Kalimantan, Ulu Sungai Mentaya, c.100 m, 30 xi 2000, Argent s.n. (holo BO, iso E), made from RBGE accession number 19942467, originally collected by Baillie G., 21 vi 1993. Figs 3, 12, 13, 14A,B.

Fig. 12.

Rigiolepis irawatiae Argent. Type, RBGE accession number 19942467. A, Branch with inflorescences (scale bar, 3 cm); B, flower (scale bar, 3 mm); C, stamens (scale bar, 1 mm); D, habit (scale bar, 3 cm); E, style with disc (scale bar, 1 mm). (Illustration: Claire Banks.)

Fig. 13.

Rigiolepis irawatiae Argent. Cultivated, RBGE accession number 19942467. (Photograph: G. Argent.)

Fig. 14.

Micrographic comparison of Rigiolepis irawatiae (A and B) and R. sulcata (C and D). Rigiolepis irawatiae, type: A, abaxial leaf surface, showing distribution of hairs; B, single dendroid hair. Rigiolepis sulcata, Paie S.45136: C, abaxial leaf surface, showing hairs; D, single multicellular hair. (Scanning electron micrographs: Frieda Christie.)

Epiphytic shrub to 2 m. Twigs with a short dense covering of simple tomentose hairs and some dendroid hairs (no brown glandular hairs), leaves lax, lenticels rounded, raised, reddish brown, prominent on older stems; lateral buds 5–7 mm long, acicular, conspicuous almost as long as petioles, scales hairy. Leaves: petiole 5–7 × c.5 mm, densely covered with both simple and dendroid hairs; blades 110–160 × 40–60 mm, ovate, narrowly ovate, elliptic to narrowly elliptic, thin, dull pale green above and below, base broadly tapering to truncate, margin entire, basal glands slightly protruding when dry at the precise junction with petiole; densely hairy with both simple and dendroid hairs below, with a similar but sparser covering above but quickly glabrescent there, 4–6 ascending veins, proximal veins arising from base and disappearing distally at about halfway up leaf, upper main lateral veins arising 20–40 mm from base and disappearing close to base of acumen, other pinnate veins from upper part of leaf less conspicuous, midvein prominent above in basal half, then impressed in a narrow channel as are lateral veins abaxially, all major veins raised, apex acuminate (to c.30 mm), extreme point acute. Inflorescence of spreading racemes from upper foliate and defoliate axils, solitary or 2 or 3 together, laxly 12- to 16-flowered, flowers pendent; basal bracts few, very small, subulate; rachis 60–110 mm, covered in curled, more or less appressed, simple hairs, dendroid hairs and occasional pale glandular hairs; bracts c.1.5 × 1 mm, ovate, concave, acute, densely covered with mainly curled simple hairs but with a few brown glandular hairs intermixed; bracteoles to c.0.9 mm long, small, subulate, variable in position but mostly subopposite and near middle of pedicel. Flowers: pedicels 7–12 × c.0.5 mm, densely covered with mainly pale long-stalked glandular hairs but with a few less conspicuous simple hairs; calyx cup-shaped, tube c.0.7 × c.1.7 mm, densely glandular hairy; lobes triangular, c.1 × c.1 mm, densely glandular hairy outside, glabrous within; corolla ovoid, slightly constricted distally, 4–4.5 × c.4 mm, pale yellow, densely glandular hairy outside, glabrous inside; lobes triangular, c.1. × 1 mm, reflexed; stamens: filaments sigmoid, c.2.5 mm, densely hairy throughout; anther cells c.0.5 mm, tubules c.0.5 mm, dorsal spurs c.0.3 mm; style glabrous, c.4 mm; disc sparsely hairy on upper side. Fruit not seen.

Description augmented from living material cultivated at Royal Botanic Garden Edinburgh.

Distribution

Indonesia: Central and West Kalimantan. Malaysia: Sarawak.

Ecology

Epiphytic in lowland dipterocarp forest at 35–200 m elevation on trees overhanging streams. Flowering: June, July, November and December, probably irregularly throughout the year.

Etymology

Named in honour of Ibu Dr Irawati, formerly director of the Kebun Raya, Bogor, who has always been helpful in promoting the taxonomy of Indonesian plants and fostering links between taxonomists and institutions.

Conservation assessment

This species has only been collected in lowland forest which is being destroyed at a very rapid rate, so it is under severe threat.

This new species is similar to Rigiolepis moultonii but differs in its extreme lowland versus montane ecology. The twigs of Rigiolepis moultonii have patently erect instead of tomentose hairs, and the leaves and inflorescences lack the dendroid hairs of this species. The basal glands protrude slightly at the junction between lamina and petiole in Rigiolepis irawatiae (not at some distance from the petiole as in R. moultonii); the flowers are densely and completely glandular hairy outside and the disc is sparsely hairy. In Rigiolepis moultonii the corolla always has some simple hairs. The new species differs from Rigiolepis bigibba and R. sulcata in the longer, laxer inflorescences (> 60 mm versus < 50 mm), the smoother leaves, which are less sulcate, the marginal leaf glands smaller, not located on a decurrent leaf base, and having a dense covering of exclusively glandular hairs on the calyx and corollas. Rigiolepis irawatiae shares with R. bigibba and R. sulcata the possession of multicellular hairs on the leaves but they are structurally different in R. irawatiae, having scabrid branches instead of smooth lamella branches (see Fig. 14).

Other than in Rhododendron, dendroid hairs are apparently unknown in Malesian Ericaceae. They have apparently not been recorded previously in Vaccinium in Malesia (Balgooy, Reference Balgooy1997). In Rigiolepis irawatiae, the hairs are of various irregular forms standing up to 220 µm high above the surfaces on which they are found. They appear under a relatively low–powered binocular microscope as a farinaceous indumentum on the lower surface of the leaves mixed with simple filamentous and glandular hairs. Each dendroid hair consists of two cells which occur side by side from a slightly bulbous base and which divide distally and branch irregularly. The two-celled nature of the stem of these hairs parallels the double nature of the stalk of dendroid Rhododendron scales in Southeast Asia. Although most similar to Rigiolepis moultonii, plants of the new species would key to Vaccinium sulcatum in Sleumer (Reference Sleumer and Van Steenis1966–1967) (now R. sulcata) on the basis of the simple hairs all over the undersurface of the leaf. As well as by the unique dendroid hairs, the new species differs from Rigiolepis sulcata by the less-bullate leaves, the much smaller basal glands and the corolla completely covered only in glandular hairs.

This species flowers regularly in cultivation in Edinburgh but fails to produce fruit, perhaps indicating that it is self-incompatible.

8. Rigiolepis jermyi Argent, sp. nov.

Most similar to Rigiolepis chaii but leaves linear, to 4 mm wide (versus 5–10 mm wide) and more strongly acute at apex. Inflorescence lacking a rachis, pedicels to 1 mm long, with impressed marginal glands 1–2 mm from the petiole. – Type: Malaysia, Sarawak, Miri District, Niah, 22 ix 1960, c.100 m elevation, Brunig S.8861 (holo SAR; iso K, L). Figs 4, 15.

Fig. 15.

Rigiolepis jermyi Argent. Type; Niah, Sarawak. A, Leaf base, showing marginal glands (scale bar, 2 mm); B, habit (scale bar, 6 cm); C, immature fruit (scale bar, 3 mm). (Illustration: Claire Banks.)

Small epiphyte. Twigs densely hairy with short semi-appressed grey hairs; lateral buds to 3 mm, much longer than subtending petioles, bud scales finely acicular. Leaves: petiole 1–1.5 × 0.5 mm, pubescent; blade 80–110 × 2–4 mm, linear, base broadly tapering, basal marginal glands distinct and impressed, 1–2 mm from petiole, apex acute; midvein impressed above, strongly raised beneath, convex with strongly recurved margins when dry with a few simple hairs beneath, without conspicuous lateral veins. Inflorescence of solitary flowers, or in 2-flowered fascicles, in leafy axils; bracteoles minute, acicular. Flowers: pedicels to 1 mm, from a few small acicular basal bracts, with semi-appressed hairs; corolla not seen. Fruit (immature) c.1.2 × 1 mm, spherical, with erect triangular calyx lobes.

Distribution

Malaysia: Sarawak, Miri Division, Niah, protected forest. Known only from the type collection.

Ecology

Epiphyte in lowland rain forest at c.100 m elevation. Flowering time not known.

Etymology

Named after Dr Anthony Clive Jermy for his contribution to Southeast Asian botany, particularly that of Sarawak, and for the inspiration he gave to many botanists.

This species is very distinct, despite not being collected in flower, with its long linear leaves with acute apex, presence of basal marginal glands and sessile flowers.

9a. Rigiolepis leptantha var. leptantha. Figs 2, 4, 16, 17.

Fig. 16.

Rigiolepis leptantha var. leptantha. RBGE accession number 19801411; Gunung Lumarku, Sabah. A, Flower (scale bar, 2 mm); B, stamens (scale bar, 1 mm); C, style with disc (scale bar, 1 mm); D, habit (scale bar, 3 cm). (Illustration: Claire Banks.)

Fig. 17.

Rigiolepis leptantha var. leptantha. Cultivated, RBGE accession number 19801411; Gunung Lumarku, Sabah. (Photograph: G. Argent.)

Small shrub to 4 m, often with a woody subglobular tuber. Twigs pendent, slender, densely puberulous with simple hairs, laxly leaved; lateral buds to c.2 mm, mostly shorter than subtending petiole, bud scales subulate and hairy. Leaves: petiole 2–5 × c.1 mm, densely puberulous; blade 45–150 × 13–40 mm, narrowly ovate to elliptic-ovate, base cuneate, margin entire, subrevolute in dry specimens, basal glands 1 or 2 on each side, small impressed on margin, c.2 mm distant from petiole, somewhat shining above, mostly dull beneath, slightly puberulent proximally beneath especially on midvein when young and with sparse, appressed glandular hairs on both sides, 3- or 5-plinerved from base, midvein deeply impressed, but prominent within depression above, obtusely raised beneath, main veins high ascending and slightly raised above, often quite obscure beneath, other shorter veins numerous, irregularly pinnate, forming a subdense, slightly prominent network above in young leaves, becoming less visible in older, dull leaves, reticulation in general inconspicuous, entire undersurface smooth, apex abruptly acuminate or subcaudate to 20 mm, often subfalcate, acute. Inflorescence a solitary raceme or 2 or 3 in a fascicle, laxly 10- to 15-flowered, faintly sweetly scented; rachis 30–55 mm, bright red, slender, with several (3–4 mm) subulate, acuminate bracts at base, densely covered with minute puberulence of very short eglandular and glandular hairs; bracts c.1.5 mm, ovate-acuminate, concave, ciliolate; bracteoles c.1 mm, subulate, inserted in lower part of pedicel. Flowers: pedicels slender to filiform, 4–8 mm at flowering, red, densely covered with minute puberulence of very short eglandular and glandular hairs, slightly accrescent in fruit; calyx tube 0.5 mm, campanulate, more densely puberulous or glandular than calyx lobes, lobes triangular, 0.6 mm, acute, ciliate; corolla 2–2.5 mm, urceolate or ovoid-subglobose, yellow or cream, puberulous in lower, glabrescent in upper half outside, lobes c.0.4 mm; stamens: filaments c.1 mm, linear, long hairy; anther cells 0.5 mm, broadly oblongoid, granular or papillose; tubules c.0.8 mm, narrow; disc pubescent, or with few hairs close to style; style 2 mm, glabrous. Fruit (immature) bright orange.

Distribution

Brunei. Indonesia: South Sumatra (possibly also in central Sumatra), Lingga Archipelago (Pulau Lingga, Pulau Sebangka), West Java, East (West Kutei) and West Kalimantan. Malaysia: Johore, Sabah and Sarawak. Singapore: Bukit Mandi.

Ecology

Generally epiphytic on tall trees, occasionally also terrestrial (in Sarawak on limestone), 10–700 m elevation. Flowering: throughout the year.

Conservation assessment

LC. This is a widespread, common species which, although it will be adversely affected by general forest clearance, will gain some protection from the fact that it often occurs on riverside trees which are valued for shade and defence against erosion.

Vaccinium flagellatifolium was reduced to a synonym of V. leptanthum by Vander Kloet (Reference Vander Kloet2005), while Sleumer distinguished V. flagellatifolium in his key by the leaves “hardly or not reticulate”, which appears to be a taxonomically trivial character. Florally and ecologically they are extremely similar, although the descriptions of the habits (“Small . . . much branched shrub” versus “shrub or small tree up to 5 m”; Sleumer, Reference Sleumer and Van Steenis1966–1967) appear to refer to very different plants. These annotations, presumably from collectors’ labels, are likely to be misleading as many collectors wrote notes long after making their collections. The Korthals specimen at K has been annotated “? part of the lectotype” by Sleumer.

Leaves: 45–100 × 15–25 mm, elliptic or ovate-elliptic, base broadly tapering to rounded, apex long acuminate or more or less abruptly acuminate, often with 2 high-ascending lateral veins arising very near base of leaf, reticulation obscure. Inflorescences with larger bracts 2–2.5 mm long and thicker (shorter) pedicels, 2–4 mm long, compared with the type.

Distribution

Indonesia: West Kalimantan. Malaysia: Sarawak.

Ecology

Reported to grow epiphytically on mangrove near Kuching also epiphytic in dipterocarp forest on alluvial soil at 50 m elevation. Flowering: not recorded.

Sleumer (Reference Sleumer1961, Reference Sleumer and Van Steenis1966–1967) distinguished Vaccinium leptanthum f. ellipticum from f. leptanthum on “the ovate-elliptic or elliptic leaves, the base broadly attenuate; pedicels thicker, (2–)3–4 mm long at anthesis”. I have modified the description with features of the Haviland specimen which is best included here, but this specimen has larger leaves, much longer attenuate leaf apices and more distinct lateral veins than the type collection. This taxon could be confused with Rigiolepis moultonii but would appear to occur in lowland habitats and lacks the distinct reticulation characteristic of the leaves of that species. Further collections and critical observations are needed to establish the taxonomic boundaries clearly. This taxon was described by J.J.Smith (Reference Smith1935) without flowers.

10. Rigiolepis linearifolia (Kloet) Argent, comb. nov. – Vaccinium linearifolium Kloet, Blumea 50: 491, f. 6 (Reference Vander Kloet2005). – Type: Malaysia, Sarawak, Miri Division (4th Div.), Kelabit, Pa Litan, Apad Runan, 1580 m, 6 v 1988, Yii Puan Ching S.55984 (holo L; iso E, K, KEP, SAN). Figs 4, 18.

Fig. 18.

Rigiolepis linearifolia (Kloet) Argent. Isotype, Kelabit, Yii Puan Ching S.55984 (E). (Scan: Robyn Drinkwater.)

Shrub. Twigs slender, pubescent; lateral buds to 5 mm long, narrowly acicular, much longer than petioles, scales sparsely patently hairy. Leaves: petiole 1 mm long, thick, hairy; blade 110–130 × 7–9 mm, linear, base broadly tapering, without lateral glands, margin entire, recurved, with long simple hairs along strongly keeled midvein underneath and scattered simple brown glandular hairs over abaxial surface, without conspicuous lateral veins, apex narrowly acute. Inflorescence of solitary, 7- to 11-flowered racemes; rachis slender, 30–50 mm long, sparsely pilose, bearing many small, caducous bracts. Flowers: pedicels c.3 mm long at flowering with a pair of bracts inserted towards base of pedicel, sparsely pilose; calyx tube small, densely hairy, calyx lobes c.1 mm long, sparingly hairy except along margins where there are also some thick brown glands; corolla globose, 3 × 4–5 mm, white, with few hairs, mostly at angles outside, lobes c.0.5 × 0.75 mm with minutely crenate margin; stamens c.2 mm long, filaments c.1 mm long, pilose; anthers c.1 mm long, oblongoid, tubules opening with an oval pore; disc glabrous; style slender, glabrous, or with a few short hairs at base, c.4 mm long. Fruit not seen.

Distribution

Malaysia: Sarawak, Miri Division, Kelabit. Known only from the type collection.

Ecology

Epiphyte, on trees in submontane mossy forest, recorded at 1580 m. Flowering: May.

The type description has been augmented from the original (Vander Kloet, Reference Vander Kloet2005) after examination of the holotype at L and isotype at E. There are no paired glands embedded on the margin of the petiole as originally described (but not illustrated), nor are there any on the margin of the leaf base. The leaves are not “± glabrous” but distinctly laxly hairy with simple hairs along the midvein underneath and with simple brown glandular hairs all over the undersurface.

Straggling or climbing shrub, to 1.5 m. Twigs at first covered with white spreading hairs; lateral buds 1–2 mm, acicular, about as long as petiole, scales with long white hairs. Leaves: petiole 1–2 × 0.5 mm, pubescent initially; blade 20–50 × 8–20 mm, 2–4 times as long as broad, narrowly to broadly ovate, base broadly attenuate to rounded, margin entire, revolute, basal marginal glands 1–2 mm from petiole, obscure, laxly hairy above but quickly glabrescent, glabrous below except for some hairs on midvein, glossy dark green above, paler and dull green beneath, slightly convex above and with a recurved margin in dry specimens, midvein strongly impressed above, prominent beneath, with 0–4 arching lateral veins, other veins from midvein irregular and numerous, forming a fine dense network or obscure, apex short or long acuminate, obtuse at extreme point. Inflorescence of one to several short, 2- to 8-flowered racemes or clusters in the upper axils; rachis 0–10 mm with several small basal bracts, shortly pubescent; bracts narrowly ovate; bracteoles narrowly ovate immediately below calyx, c.2 mm, ribbed lengthwise when dry, shortly pubescent. Flowers: pedicels to 1 mm; calyx tube obconical, c.1 mm, lobes c.2 × 0.5 mm, narrowly ovate, distinctly ribbed lengthwise when dry, shortly pubescent, terminating in a gland which is brown when dry; corolla c.2.5 × 2 mm, subglobose-urceolate, slightly 5-angled, white, shortly pubescent outside; stamens: filaments linear, sparsely hairy, 0.5–0.7 mm; anthers cells oblongoid, c.0.5 mm, tubules c.0.5 mm, cleft nearly to base; disc hairy; style papillose, c.2 mm long. Fruit 6–7 mm in diameter, hairy, with persistent calyx lobes.

Chromosome number

2n = 24 (Atkinson et al., Reference Atkinson, Jong and Argent1995).

Distribution

Malaysia: Sarawak.

Ecology

On limestone ridge, 450–900 m. Flowering: August.

Conservation assessment

Although of unknown size, populations of this species are likely to be vulnerable to deforestation and El Niño droughts because of their lowland habitat.

11b. Rigiolepis lobbii var. lanceifolia (Ridl.) Argent, stat. nov. – Rigiolepis lanceifolia Ridl., Bull. Misc. Inform. Kew 106 (1922). – Vaccinium lanceifolium (Ridl.) Sleumer, Biol. Jahrb. Syst. 71: 168 (Reference Sleumer1940) (‘lancifolium’). – Type: Malaysia, Sarawak, near Quop, without date, Haviland 619 (lecto K, designated by Sleumer, Reference Sleumer1961, p. 16; isolecto SAR). Figs 3, 19, 20.

Fig. 19.

Rigiolepis lobbii var. lanceifolia (Ridl.) Argent. RBGE accession number 19820748; Gunung Bungo Range, Sarawak. A, Flower with bracts (scale bar, 2 mm); B, stamens and style with disc (scale, bar, 1 mm); C, bare branch with inflorescences (scale bar, 1 cm); D, habit (scale bar, 5 cm). (Illustration: Claire Banks.)

Fig. 20.

Rigiolepis lobbii var. lanceifolia (Ridl.) Argent. Cultivated plant with ripening fruit and flowers, RBGE accession number 19820748; Gunung Bungo Range, Sarawak. (Photograph: G. Argent.)

Spreading shrub, to 1.5 m. Twigs at first covered with white spreading hairs; lateral buds 1–2 mm, acicular, about as long as petiole, scales with long white hairs. Leaves: petiole 1–2 × 0.5 mm, pubescent initially; blade 50–80 × 6–15 mm, 5–8 times as long as broad, lanceolate to narrowly elliptic, base broadly attenuate to rounded; margin entire, weakly to strongly recurved, basal glands marginal, 1–2 mm from petiole, obscure, laxly hairy above but quickly glabrescent, glabrous below except for some hairs on midvein, glossy dark green above, paler and dull green beneath, slightly convex above, midvein strongly impressed above, prominent beneath, with one pair of intramarginal veins which are often obscure, and smaller pinnate veins from upper part of leaf. Inflorescence as for var. lobbii.

Chromosome number

2n = 24 (Atkinson et al., Reference Atkinson, Jong and Argent1995).

Distribution

Malaysia: Sarawak.

Specimens cultivated in Edinburgh retain their distinctive leaf shapes, var. lanceifolia having much larger, especially longer leaves than var. lobbii. Both taxa were collected on Gunung Buri and are are nearly devoid of glandular hairs.

Shrub to 5 m. Twigs minutely, shortly hairy; lateral buds with acicular scales as long as subtending petioles but eroding to short tufts. Leaves: petiole 4–10 × c.2 mm, grooved above, pubescent; blade 200–300 × 60–100 mm, elliptic-ovate, base rounded, margin entire, more or less flat, basal glands impressed on margin at some distance from petiole, without hairs except for midvein and main lateral veins which are shortly pubescent basally, laxly covered with simple brown glandular hairs beneath, with 4–6 strongly high-arching veins and with 7 or 8 distinct pinnate veins from each side, which are a little decurrent along midvein, main veins prominent in a pronounced depression above, strongly raised beneath, apex long-acuminate, the narrow acumen (30–50 mm) obtuse, reticulation conspicuous and dense, slightly raised on both sides. Inflorescence a solitary raceme or commonly a fascicle of racemes, 10- to 30-flowered; rachis 20–60 mm, puberulous; bracts 1.5 mm long, ovate, concave; bracteoles not seen. Flowers: pedicels 2–4 mm; calyx small, pubescent, lobes inflexed; corolla urceolate, 2–3 mm long, white or cream; stamens c.2 mm long, filaments not seen; style glabrous, 2–3 mm long; disc glabrous. Fruit (immature) c.3 mm in diameter, subglobose, laxly pubescent, calyx lobes inflexed.

Distribution

Indonesia: Central Kalimantan, Mount Batu Lesong.

Ecology

Epiphytic on tree, summit vegetation. Flowering: January.

I have taken the very brief floral description from Vander Kloet (Reference Vander Kloet2005). Nooteboom & Chai 1692, collected on 25 iii 1970 at Bintulu Division, Bario, Kelabit Highlands, is tentatively placed here. It has no corollas but small (c.1 mm) styles and both very large (to 170 × 80 mm) and small leaves (c.70 × 20 mm), both with infloresences in their axils. This specimen keys to Rigiolepis macrophylla with the large leaves, but if only the small leaves were present on the specimen, it would be similar to R. minimiflora although it has a glabrous disc. It might represent an undescribed species.

Shrub or climber c.1.5 m. Twigs shortly pubescent when young; lateral buds 2–2.5 mm, as long as or slightly longer than petioles. Leaves: petiole 1–2 mm, pubescent; blade 30–60 × 10–20 mm, narrowly ovate to ovate, base rounded; margin entire, slightly revolute when dry, basal pair of glands small, c.1.5 mm distant from petiole, puberulous at midvein and ciliate initially, some pubescence remaining at base of older leaves, laxly glandular punctate beneath, glossy on both sides, with 2 or 3 pairs of high-ascending main veins, with several shorter, less conspicuous veins from upper part of midvein, inconspicuous above except midvein and lowest pair of veins which are slightly raised; reticulation dense and slightly raised beneath; apex subcaudate-acuminate, extreme tip obtuse or acute. Inflorescence: lax, axillary racemes, solitary, many-flowered; rachis 25–40 mm, slender, shortly pubescent and with a few stalked glands; bracts 0.7 mm, subulate; bracteoles subulate, inserted close below calyx. Flowers: pedicels 1.5–2 × 0.5 mm, shortly pubescent, with simple hairs and a few brown glandular hairs; calyx tube 0.8 mm, campanulate, densely hairy, lobes 1–2 mm, deltoid, sparsely hairy, acute; corolla c.3 mm, urceolate, contracted distally, whitish green or red, laxly covered with very short, rufescent, glandular hairs at base, glabrous apically; stamens: filaments c.1 mm, hairy; anther cells c.1 mm, with 2 long dorsal spurs; tubules 1 mm; disc shortly hairy; style 3 mm, glabrous. Fruit: not seen.

Distribution

Malaysia: Sarawak, Ulu Sungai Koyan, Bukit Dulit.

Ecology

Mossy forest, c.1000 m. Flowering: March to August.

Vander Kloet (Reference Vander Kloet2005) says “except for the presence of a rachis and multiple flowers, similar to Vaccinium uniflorum”. This may well be so but there are far too few collections to evaluate this species properly. Vander Kloet (Reference Vander Kloet2005) expands the leaf size from Sleumer's (Reference Sleumer and Van Steenis1966–1967) description (30–70 × 10–20 mm versus 30–40 × 10–15 mm). Lee S.54556 has even larger leaves, to 140 × 30 mm.

Distribution

Indonesia: East Kalimantan. Malaysia: Sabah and Sarawak.

Ecology

Epiphytic in submontane forest at c.600–1250 m. Flowering: November.

Conservation assessment

LC. Widespread with some populations in protected forest (Mount Murud), but the situation at the type locality is unknown.

This is still a somewhat problematic species taxonomically. J.J.Smith (Reference Smith1935) did not describe the flowers of Rigiolepis dipladenium, but Sleumer (Reference Sleumer and Van Steenis1966–1967) added a partial description from what he described as “rests adhering to a young fruit”. I have included Vaccinium dipladenium as a synonym of Rigiolepis moultonii because the key characters used by Sleumer (Reference Sleumer and Van Steenis1966–1967) to distinguish these species are the reticulation of the leaf veins, a character which is somewhat subjective, and a difference in the pedicel length (2–4 mm in V. moultonii versus 4–7 mm in V. dipladenium). Because the pedicel elongates as the fruit develops and Vaccinium dipladenium was described from “young fruit”, the difference in length would appear to be insignificant; neither would the apparent difference that the racemes of V. dipladenium were described without glandular hairs. I have thus also included Vaccinium dipladenium in as a synonym. The montane ecology and other features agree well.

Rigiolepis moultonii has been confused with R. salicifolia. Rigiolepis moultonii mostly has broader, ovate leaves with conspicuously high-arching basal veins and slender pedicels while R. salicifolia has more narrowly ovate or elliptic leaves with dominant pinnate venation, the basal veins being much less conspicuous. The pedicels are also shorter and thicker in that species. As conceived here, Rigiolepis moultonii is a widespread but variable species. A better evaluation awaits new collections from the type locality and other localities.

14b. Rigiolepis moultonii subsp. muluensis Argent, subsp. nov.

Differing from subspecies moultonii in the broader leaves with longer, broader petioles, the larger flowers, style longer 5–5.5 mm (versus 3.5–4 mm), sometimes hairy near the base and the hairy disc. From subspecies murudensis in the smoother leaves, the veins only weakly raised abaxially and the calyx tube with simple hairs more prominent than the small glandular hairs. – Type: Sarawak, Miri Division, Gunung Mulu National Park, 10 iii 2017, Argent s.n. (RBGE accession number 19773554). Cultivated material Kerby, 297 (holo SAR, iso E). Figs 3, 21–23.

Fig. 21.

Rigiolepis moultonii subsp. muluensis Argent. RBGE accession number 19773554. A, Flower (scale bar, 5 mm); B, stamens (scale bar, 2 mm); C, style with disc (scale bar, 2 mm); D, habit (scale bar, 5 cm). (Illustration: Claire Banks).

Fig. 22.

Rigiolepis moultonii subsp. muluensis Argent. Flowering plant, RBGE accession number 19773554; Gunung Mulu, Sarawak. (Photograph: G. Argent.)

Fig. 23.

Rigiolepis moultonii subsp. muluensis Argent. Plant with ripe and immature fruit, RBGE accession number 19773554; Gunung Mulu, Sarawak. (Photograph: G. Argent.)

Distribution

Malaysia: Sarawak, known from the type locality, Mount Mulu.

Ecology

Montane mossy forest epiphytic and terrestrial on peaty ground, 1800–2300 m elevation. Flowering: November, in cultivation February to March.

Conservation assessment

LC. There is a good population on Mount Mulu which is in a well-protected national park.

The most significant difference between this subspecies and both the other subspecies is that the disc is densely hairy, not glabrous and the larger flowers reflected by the longer styles. There are small vegetative differences, the petiole being mostly longer and much broader and the reticulation being much less conspicuous.

Distribution

Malaysia: Sarawak, Mount Murud.

Ecology

Montane forest and shrubberies.

This is a distinctive subspecies in its narrow sulcate leaves, mostly from Mount Murud.

Shrub to c.3 m. Twigs with both simple and glandular hairs, laxly leaved; lateral buds 4–8 mm long, narrow with subulate scales which are almost as long as petiole. Leaves: petiole 5–10 × 2–3 mm rugose, glabrous, or hairy; blade 180–300 × 50–100 mm, large, narrowly ovate to elliptic, base broadly tapering to rounded, margin entire, shortly revolute, basal glands distinct but not protruding beyond leaf margin and inserted 1–3 mm from petiole, puberulous in basal part of midvein, otherwise glabrous except for some sparse glandular hairs abaxially, with 4–6 high-arching lateral veins from and from a little above base, midvein and inner two pairs of veins prominent above within a shallow depression, strongly prominent beneath, outer pair of veins much less distinct, numerous shorter veins pinnate from distal part of midvein, irregular, slightly impressed above, a little raised beneath, veins forming a dense, fine prominent network on both sides; apex subcaudate to long acuminate, acute. Inflorescence of solitary racemes, laxly c.10-flowered, with several subulate, very acute bracts (4–6 mm) at base, which also occur in sterile axils; rachis slender, 20–60 mm, puberulous and minutely glandular hairy; bracts 1.5–2 mm, ovate-acuminate; bracteoles minute, subulate. Flowers: pedicels 2–3 mm, slender, indumentum as on rachis; calyx laxly to densely hairy, with some glandular hairs, tube semiglobose, c.0.5 mm, lobes ovate acuminate, subacute, c.1 mm; corolla c.3 mm, ovoid urceolate, thin, green or white?, laxly covered with short muriculate glandular, and sometimes simple, white hairs; stamens: filaments 0.5 mm long, hairy; anther cells broad oblongoid, papillose, c.0.7 mm; tubules 0.7 mm, disc laxly pubescent; style 2.5 mm, glabrous. Fruit: not seen.

Distribution

Malaysia: Sarawak, Gat, Upper Rejang River, Kapit Division, Belaga District, Bukit Dema.

Ecology

On large trees overhanging rivers at 900–1100 m elevation. Flowering: July to September.

The bracteoles vary in position within an inflorescence from alternate in the proximal half of the rachis, to alternate about the middle of the rachis or subopposite at the distal end of the rachis (the uppermost flower) (Burtt 11340). Vander Kloet (Reference Vander Kloet2005) reduces this species to Vaccinium acuminatissimum Miq., but based on the small non-protruding glands, it would appear to be better maintained or possibly amalgamated with Rigiolepis macrophylla.

Shrub straggling or climbing to 1.5 m. Twigs densely white pilose, bud scales long and pointed, pilose; lateral buds to 3 mm, longer than adjacent petiole, densely white hairy. Leaves: petiole 1–2 × c.1 mm, pilose initially; blade 50–100 × 15–20 mm, very narrowly ovate, base broadly tapering to rounded, margin entire, narrowly revolute, basal glands strictly marginal, 1.5–2 mm from petiole, apex long acuminate and narrowly acute, midvein impressed above throughout its length, strongly raised below and there densely hairy, mostly with 2 weak arching veins which disappear just beyond mid-leaf, lamina laxly pilose both above and below. Inflorescence: 1–3 racemes bearing 8–15 flowers; rachis slender, 40–60 mm long, densely white pilose (hairs to 0.6 mm); bracts 4–5 mm, ovate, concave, exceeding the pedicels, densely pilose; bracteoles narrowly ovate, c.2 mm, pilose. Flowers: pedicels 1–2 mm long at flowering, pilose; calyx tube 0.7–0.8 mm, lobes c.3 mm long, ribbed when dry, terminating in a brown gland; corolla 2.8 × 2.5 mm, broadly ovoid, hairy outside, glabrous inside; lobes 0.8 mm, broadly triangular, densely hairy outside; stamens: filaments 1 mm, glabrous, anthers cells c.1 mm long; tubules c.0.7 mm long, opening with an elongated slit; style 3.2 mm, glabrous, narrowly club-shaped; disc hairy. Fruit: not seen.

Distribution

Malaysia: Sarawak, Kuching Division, Berumput, Mount Poi Range, Mount Serapi (Matang).

Ecology

Commonly epiphytic in low open forest, and rocky summits, 1000–1700 m elevation. Flowering: May to August.

This species represents the more slender-leaved forms of Rigiolepis with costate bracts and calyx lobes. It has dense long white hairs and long inflorescences. Plants of this group cultivated in Edinburgh over many years show leaf size and inflorescence length to be genetically fixed and remain constant under identical growing conditions. There are considerable discrepancies between the protologue of Vaccinium crinigerum (Vander Kloet, Reference Vander Kloet2005) and the holotype specimen in Leiden. The pedicels are shorter than in the type description and the position of the bracteoles is not “immediately below the calyx” but extremely variable in position as observed in other species. The calyx lobes do not end in a pair of bristles but mostly terminate in a prominent brown gland (as in the related Rigiolepis lobbii). The corolla is described by Vander Kloet as red but the colour is not recorded on the field label and there is no mention of the hairs in the type description which are a prominent feature on the outside of the corolla (well described in the type of Rigiolepis poiana by J.J.Smith, Reference Smith1935), although these are illustrated by Vander Kloet. Vaccinium crinigerum is completely consistent with Rigiolepis poiana as described by J.J. Smith and thus reduced to synonymy.

I have followed Sleumer (Reference Sleumer1961) in using Foxworthy 395 as the type despite the fact that it is almost certainly destroyed in PNH. Sleumer clearly designates the Clemens specimens that he has not seen but by implication he has seen Clemens 20043 n.v. and 22589 n.v., both cited in the protologue and from which a lectotype could be selected if it can indeed be demonstrated that the Foxworthy specimen has been destroyed.

Shrub to 4 m. Twigs slender, shortly pubescent; lateral buds to 2 mm long, densely hairy, mostly shorter than subtending petiole, bracts acicular from a broad base. Leaves: petiole 1.5–5 × 1–2 mm, somewhat flattened, grooved above; blade 48–160 × 8–45 mm, ovate-elliptic to narrowly ovate-elliptic, base broadly attenuate to rounded, slightly decurrent at upper part of petiole, margin entire, broadly revolute when dry, basal glands marginal, distinct, impressed, at junction of lamina and petiole or up to 2 mm distant from petiole, glabrescent except for some hairs on proximal part of midvein, very laxly covered with appressed glandular hairs abaxially, with 4–6 high-arching main veins and with 2–4 pairs of pinnate nerves from midvein, which are as distinct as basal ones, these a little decurrent along midvein, steeply high ascending, anastomosing; main veins slightly raised within a shallow depression above, sharply prominent beneath, apex short- to long-acuminate or subcaudate, acute or nearly so, reticulation dense, finely prominent on both sides. Inflorescence: solitary racemes or several in a fascicle, drooping laxly, 7- to 18-flowered, flowers described as fragrant; rachis slender, 25–50 mm, finely pubescent and glandular muriculate; bracts c.2.5 × 1 mm, oblong, ciliolate; bracteoles c.1 mm long, narrowly triangular, fringed with glands. Flowers: pedicels 1.5–6 mm, variable in thickness, finely pubescent and glandular muriculate; calyx tube c.1 mm, obconical, becoming subglobose, finely pubescent and glandular hairy; lobes 1 mm long, triangular, acute, pubescent; corolla c.4 × 3 mm subglobose-urceolate, 5-angular, light green becoming red, glabrous or laxly glandular hairy, lobes c.0.6 mm; filaments 1.5–2 mm linear, wavy, patently hairy; stamens c.3 mm, anthers 1.1 mm long, oblongoid, tubules 0.6 mm; disc minutely hairy or nearly glabrous; style 3.5 mm, glabrous. Fruit spherical, c.5 mm in diameter, passing through yellow to red.

Distribution

Indonesia: East Kalimantan, West Kutei, Mount Kemul and Mount Palimasan. Malaysia: Sarawak, Miri Division, Mount Mulu; Batu Lawi.

Ecology

Mostly epiphytic in submontane mossy forest, also on white acid sands and on rocky mountain ridges, 800–1800 m. Flowering: June to October.

Conservation assessment

LC. Common and widespread.

This species is usually distinctive in its narrowly ovate leaves, prominent pinnate venation and relatively large flowers on robust rachis and pedicels. It is sometimes confused with Rigiolepis moultonii (see notes under that species).

Semi-epiphytic shrub. Twigs slender, densely floccose hairy; lateral buds 3–5 mm long, acicular, hairy, as long as petioles. Leaves: petiole 3–5 × c.1.25 mm, densely floccose hairy; blade elliptic to subovate, 120–140 × 30–50 mm, base broadly tapering, margin entire, narrowly revolute, basal marginal glands small, impressed, 6–8 mm from base of blade, apex acute from a 10–20 mm acumen; glabrous above or with a few hairs on main veins initially, sparsely glandular hairy below; main veins 3, high ascending, laterals disappearing at base of acumen, deeply impressed above and strongly raised below, with distinct scalariform pattern of tertiary veins visible beneath; Inflorescence of very short lateral racemes; rachis 1–4 mm long, hairy, 1- to 3-flowered; bracts c.3 mm, narrowly ovate, distinctly ribbed when dry; bracteoles c.1.5 mm, ribbed. Flowers: pedicels c.1 mm, hairy; calyx tube c.3 mm, densely hairy, lobes c.3 mm long, distinctly ribbed lengthwise, fringed with both simple and glandular hairs; corolla, style and stamens not seen. Fruit hairy, orange (immature), 4–5 mm in diameter.

Distribution

Indonesia: West Kalimantan. Known only from the type collection.

Ecology

Not described but collected at only 10 m elevation. Flowering: not known.

Conservation assessment

Likely to be vulnerable due to large-scale forest destruction at low elevation, even though the type locality is in a national park.

This species is superficially similar to Rigiolepis tenax but differs by a distinctive scalariform tertiary leaf venation, marginal glands on the leaves 6–8 mm from the petiole (versus 2–3 mm) and the calyx lobes c.3 mm long with distinct multiple ribs (versus c.1 mm with usually a single prominent midrib). It is probably most closely related to Rigiolepis borneensis, which differs in having a much stronger inflorescence with more flowers and a rachis at least 10 mm long.

Epiphytic shrub to 5 m. Twigs pubescent, covered in tomentose grey hairs and yellowish or brownish dendroid hairs; lateral buds 3–5 mm, long, to just over half the length of petiole, densely covered in short hairs. Leaves: petiole 5–8 × 2–3 mm, thick, pubescent; blade 100–280 × 35–65 mm; ovate, narrowly ovate or elliptic, base rounded to broadly tapering, apex caudate-acuminate (20–30 mm), margin entire, revolute, with a small decurrent auricle at base on which are 2 prominent marginal glands, remaining laxly hairy with simple hairs all over the underside, with or without glandular hairs and with multicellular hairs, leathery, mostly sulcate with distinct reticulation at least on upper side, initially hairy with simple hairs especially on main veins beneath, also with multicellular hairs that give a yellowish scurfy appearance, quickly glabrescent above, with usually 5 main veins, laterals arching upwards to base of acumen, occasionally subpinnate, arching veins disappearing at about mid-leaf, with much smaller veins in upper part of leaf. Inflorescence of 2–4 racemes in axillary fascicles, slender, subdensely covered with simple pale hairs and much shorter, darker glandular hairs; rachis slender, 30–50 mm long; bracts c.1.5–2 mm long; bracteoles subulate, mostly around middle of pedicel. Flowers: pedicels slender 5–10 mm long; calyx tube 0.5 mm, broadly obconical, covered with short glandular hairs and longer simple hairs, lobes 1–1.5 mm long, triangular; corolla 4–4.5 × 2.5–3 mm, urceolate, slightly contracted distally, pale yellow, laxly hairy with simple hairs especially at longitudinal angles, glabrous between, lobes c.0.5 mm; filaments 2.5 mm, linear, densely hairy; anther cells 1 mm, oblongoid, echinulate, dorsal spurs c.0.5 mm, tubules c.1 mm; disc hairy; style 4 mm, glabrous. Fruit c.4 mm, hairy.

Distribution

Indonesia: West Kalimantan. Malaysia: Sarawak.

Ecology

In primary and secondary forest at low to moderate elevations, also on limestone. Flowering: June.

There appears to be considerable variation in the indumentum on the underside of the leaves. The type collection has branched hairs with flat lamellar extremities from a double-celled base. Some collections have a low, densely glandular surface. More observations are needed to see if the variations in indumentum are taxonomically significant.

Fig. 24.

Rigiolepis tenax (Argent) Argent. Type, Burtt & Martin B4765. A, Habit; B, flower; C, pistil with calyx; D, stamens. (Illustration: Dorothy Brunton.)

Epiphytic shrub, to 2 m, climbing by adventitious roots. Twigs pubescent with short glandular hairs and less dense, longer simple hairs; lateral buds narrowly pyramidal, scales acicular about half as long as petiole, often eroding to much shorter points. Leaves: petiole c.5 × 2 mm, for the most part winged with decurrent lamina margins; blade 130–200 × 30–95 mm; ovate-acuminate to elliptic, base rounded, decurrent at very base in narrow wings down petiole, margin entire, very narrowly revolute, glabrous above, sparsely glandular hairy below and with a few non-glandular hairs on veins, apex acute, or acuminate to c.20 mm, main veins 3–5, high ascending, slightly raised in lower half above, strongly raised below, basal glands small, on rounded margin, 2–3 mm from petiole. Inflorescence of solitary flowers or two to several in a fascicle or short raceme; rachis 0–2 mm long; bracts c.0.5 mm, small; bracteoles 0.8 mm, narrowly triangular, fringed with simple hairs. Flowers: pedicels 2–3 mm long with semi-appressed or patent, whitish and a few brown glandular hairs; calyx tube c.0.7 mm, lobes c.0.5 mm, glandular hairy or with simple white hairs, costate with a single midvein when dry; corolla urceolate, somewhat 5-angled, c.2 mm long with scattered glandular hairs outside, greenish-cream; stamens: filaments c.0.5 mm, glabrous except for a tuft of hairs adaxially at base; anthers c.1.25 mm long, glabrous but slightly papillose at base; disc pilose with erect hairs; style 1.5 mm long, glabrous but slightly papillose at base. Fruit (immature) spherical, orange.

Distribution

Brunei: Belait, Ulu Sungai Ingei. Indonesia: West Kalimantan. Malaysia: Sarawak, Miri Division, Mount Mulu National Park, Melinau terraces.

Ecology

Heath and peat swamp forest, 25–202 m elevation. Flowering: August to September, probably at other times.

Conservation assessment

LC. Widespread in occurrence with several records from well-protected areas.

This species was overlooked by Vander Kloet (Reference Vander Kloet2005) and therefore not considered in his account. It is superficially similar to his Vaccinium suberosum; for differences, see under this species.

Argent & Sinclair 8219, 4 iii 1982, collected in Sarawak, Kuching Division, Gunung Bunga Range, Seropak village, is a sterile collection which may be referable to this species. The ecology of this specimen (8219) is somewhat different from that described above for the species, being from a summit ridge in pole forest at c.800 m. The leaves are very similar to those of Rigiolepis suberosum, where the inflorescences are described as short but are not present on the holotype.

Distribution

Indonesia: Kalimantan, Central, Amai Ambit. Malaysia: Sarawak, Ulu Sungai Mayeng, Kakus.

Ecology

Epiphyte in submontane heath forest or terrestrial on limestone, 600–900 m. Flowering: May, August, November, probably intermittently throughout the year.

Fig. 25.

Rigiolepis uniflora var. monantha (Ridl.) Argent. RBGE accession number 19773328; Gunung Mulu, Sarawak. A, Flower (scale bar, 2 mm); B, stamens (scale bar, 1 mm); C, style with disc (scale bar, 1 mm); D, habit (scale bar, 3 cm). (Illustration: Claire Banks.)

Fig. 26.

Rigiolepis uniflora var. monantha (Ridl.) Argent. Plant with flowers and fruit, in cultivation, RBGE accession number 19773329; Gunung Mulu, Sarawak. (Photograph: G. Argent.)

Flowers: with the calyx and corolla glabrous or with sparse glandular hairs only, otherwise as in the nominate variety.

Distribution

Malaysia: Sarawak, Miri Division, Niah; Baram District, Mount Trekan, Mount Mulu.

Ecology

Epiphytic in lowland to submontane heath forest or sometimes terrestrial on limestone, 100–1100 m elevation. Flowering: throughout the year.

Conservation assessment

LC. Widespread with several localities in well-protected areas.

I have followed Vander Kloet (Reference Vander Kloet2005) in uniting Vaccinium monanthum Ridl. with Vaccinium uniflorum J.J.Sm. although retaining them as varieties. They are often common forest epiphytes but there is too little information on the variation of the indumentum to be confident of their status.

Distribution

Malaysia: Sarawak.

Ecology

Epiphytic on a tree, elevation unknown. Flowering: May to October, probably throughout the year.

This variety is distinguished from the other varieties of Rigiolepis uniflora by its more slender leaves, long pedicels and occasional short rachis surmounted by a solitary flower. The flowers are covered with glandular hairs, as is common in Rigiolepis uniflora var. monantha. Leaf size and shape are most similar to Rigiolepis lobbii var. lobbii but the indumentum of abundant glandular hairs intermixed with simple hairs on the stems is quite different and R. uniflora var. pedicellata lacks the large costate bracts and calyx lobes of R. lobbii.

Fig. 27.

Rigiolepis uroglossa var. uroglossa. RBGE accession number 20080419A; Gunung Alab, Sabah. A, Flower (scale bar, 3 mm); B, stamens (scale bar, 1.5 mm); C, style with disc (scale bar, 1.5 mm); D, habit (scale bar, 3 cm). (Illustration: Claire Banks.)

Fig. 28.

Rigiolepis uroglossa var. uroglossa. Flowering plant, Gunung Alab, Sabah. (Photograph: G. Argent.)

22a. Rigiolepis uroglossa var. uroglossa. Figs 3, 27, 28.

Epiphytic or semiclimbing shrub or small tree, to 5 m, often with a globose tuber. Twigs densely grey-hairy with subpatent simple hairs and a few brown-tipped glandular hairs, leaves densely to laxly arranged; lateral buds 3–4 × 1.5–2 mm, outer scales finely subulate, much longer than subtending petioles. Leaves: petiole 1.5–2.5 × 1 mm, densely hairy; blade 30–65 × 14–22 mm, ovate to narrowly ovate, base rounded, margin entire, narrowly revolute, ciliate initially; basal pair of marginal glands small, c.2–3 mm from petiole but not always present, pubescent, glabrescent with age although retaining some indumentum at base and lower part of main veins, subdensely to laxly covered with fine appressed, club-shaped, glandular hairs on both sides, often persisting beneath, finally glabrescent, flat, with 2 or 3 high-arching veins from base and from somewhat above base, several other upper veins irregular, and spreading, less distinctly raised than mid- and basal veins, reticulation dense and minutely prominent on both sides, apex acuminate to subcaudate, extreme tip obtuse. Inflorescence: of solitary, secund, racemes, densely 10- to 20-flowered; rachis slender, 40–60 mm, subdensely covered with white, short hairs mixed with slender glandular hairs, with several subulate, stipule-like bracts (2–4 mm) at base; bracts ovate, subulate, nearly as long as pedicel; bracteoles subulate, inserted near base or in lower half of pedicel, 1–1.5 mm. Flowers: pedicels slender, 2–4 mm, with hairs similar to those on rachis; calyx tube, c.1 mm, campanulate, densely white, hairy and with brown-tipped glandular hairs, lobes 0.8 mm ovate-triangular, more sparsely hairy than tube; corolla 4–4.5 mm, urceolate, slightly 5-angular, greenish to cream or yellow turning pink with age, laxly covered with subclavate, glandular and rarely some simple hairs at angles of tube, lobes 0.5 mm; stamens: filaments 1.5 mm, slightly wavy, linear, densely hairy; anther cells 0.8 mm, granular, tubules c.1.5 mm; disc densely pubescent; style 3.5 mm, glabrous. Fruit c.8 mm in diameter, initially green, turning orange and finally red when fully ripe.

Distribution

Brunei: Mount Pagon Ridge. Malaysia: Sabah, Mount Kinabalu, Crocker Range, Mount Alab; Sarawak.

Ecology

Often common as an epiphyte in montane mossy forest, occasionally terrestrial in open disturbed sites, 900–1830 m elevation. Flowering: January to December.

Conservation assessment

LC. A widespread species growing in relatively undisturbed habitats, several of which have protected status.

See discussion of varietal status below. Plants of this species in flower on Mount Alab were thronged by small bees.

In the protologue, Sleumer (Reference Sleumer1940) cited Clemens 29854 at B and BM as the type collection without indicating which was holotype. Later, he cited only the duplicate at BM as type (Sleumer, Reference Sleumer1961), so this is an effective lectotypification.

Differing from the type variety in the laxer disposition of broader leaves (to 30 mm wide), the frequent lack of marginal leaf glands and a southern distribution.

Distribution

Brunei. Malaysia: Sabah and Sarawak.

Ecology

Epiphytic in submontane, mossy forest, 950–1800 m elevation. Flowering: January to December.

Conservation assessment

LC. Widespread and well protected in conservation areas.

I have included Vaccinium tenerellum within Rigiolepis uroglossa. Vander Kloet (Reference Vander Kloet2005) suggested that they should be amalgamated. The distinction Sleumer makes in his key (Reference Sleumer and Van Steenis1966–1967) of “calyx densely hairy vs. calyx sparsely glandular, epilose” breaks down. There is great variation in the indumentum of the calyx from densely covered with simple white hairs, as in the classic locality on Kinabalu, to glandular hairs, as on the type of Vaccinium tenerellum from Brunei, and various mixtures of simple and glandular hairs commonly occur. In the northern part of the distribution (Sabah), the leaves have conspicuous marginal glands and are densely overlapping. Further south, in Sarawak, the plants are mostly without or have very inconspicuous marginal glands, and have a lax arrangement of broader leaves, but the distribution of these forms is not sufficiently consistent to be used to separate Vaccinium tenerellum from V. uroglossum at the species level.

A specimen from Mount Besar, South Kalimantan (Kato & Mogea 3459 BO!) is in fruit only. It has impressed basal glands some distance from the petiole and is similar to Rigiolepis uroglossa but a very long way from other locations of this species which are otherwise all to the northwest of Borneo. A specimen from the Hose Mountains (Ashton S.19097) has both simple and glandular hairs on both calyx and corolla, and the glandular hairs are on long stalks, which gives a very distinctive appearance to this collection. The calyx lobes are also more distinctly ribbed than is usual in this species, leading Sleumer to comment on the sheet “prob. n.sp.” which may well be true.

23. Rigiolepis winkleri Argent, sp. nov.

Leaves narrowly ovate, 90–135 × 15–40 mm. Similar to Rigiolepis moultonii but differing in its smaller, narrower leaves; by having long-stalked glandular hairs which overtop dense simple hairs on the inflorescence, these also being found on young leaves, and a hairy disc. – Type: Indonesia, West Kalimantan, Bukit Raja, c.1400 m, 21 xii 1924, H. Winkler s.n. (holo HBG, iso L). Fig. 29.

Fig. 29.

Rigiolepis winkleri Argent. I. Paie S.26588, Bukit Salong, Sarawak. (Scan: Robyn Drinkwater.)

Epiphytic shrub. Twigs with a short dense indumentum of simple grey hairs which are overtopped by much longer brown glandular hairs, leaves complanately arranged; lateral buds 2–3 mm long, acicular, almost as long as petioles, scales covered with simple and long glandular hairs. Leaves: petiole 2–4 × c.1 mm, densely covered with both simple and glandular hairs; blade 80–135 × 15–40 mm, narrowly ovate, base broadly tapering to rounded, margin entire, flat to narrowly recurved, basal glands impressed, 2–3 mm distant from petiole, apex long acuminate (to 4 cm), narrowly acute, thin papery in texture, with simple glandular hairs below and with simple and long-stalked glandular hairs alongside mid- and main veins, glabrescent above but with a few hairs persisting on and near main veins, mostly tripliveined, these arising from just above base, other veins much less conspicuous, a pair of weaker veins sometimes from near base and a few short pinnate veins distally; midvein prominent above within a groove, distally totally impressed as are main lateral veins, all major veins prominently raised below, reticulation distinct, especially below. Inflorescence: of spreading racemes from upper foliate axils, solitary, laxly 5- to 6-flowered; basal bracts few, very small, subulate; rachis 20–40 mm, densely hairy with short, patent simple white hairs and much longer, more conspicuous, stalked brown glands; bracts c.1.5 × c.0.75 mm, ovate, acuminate, covered with simple and long-stalked glandular hairs; bracteoles c.0.5 × 0.25 mm, minute, ovate, attached about halfway along pedicel. Flowers: pedicels 4–6 × c.0.5 mm, densely covered with short simple hairs and much longer brown glandular hairs; calyx tube to c.1 mm, hemispherical, densely glandular hairy, lobes 1.5–2 × c.1 mm, weakly costate or smooth, broadly triangular, densely glandular hairy outside, more sparsely hairy within; corolla 3.5 × 2 mm, barrel-shaped, with a few simple hairs outside; stamens to 2.5 mm, filaments 0.7 mm with long hairs throughout, anthers cells 0.8 mm, echinulate, tubules 1 mm; style 2.7 mm, glabrous; disc densely hairy with erect hairs on upper side. Fruit (described from I. Paie S.26588) orange (immature), spherical c.5 mm in diameter.

Distribution

Indonesia: South Kalimantan. Malaysia: Sarawak.

Ecology

Epiphytic shrub 4 m high on host tree, in montane forest on hillside at c.(50–)800–1400 m elevation. Flowering: December.

Etymology

Named after Hans Winkler (1877–1945), collector of the type specimen and eminent German botanist.

Distinctive with narrower leaves than is usual in Rigiolepis moultonii and the possession of both simple and long-stalked glandular hairs on stems, young leaves and especially on the inflorescence. Argent & Coppins 1200a, from Gunung Mulu National Park, 8 v 1978, agrees well but has no inflorescence and cannot be determined with certainty. Argent & Saridan 9323 from Central Kalimantan, Kotawaringin Timur, Sungai Mentaya, Km 54 N of Sangai, 31 i 1993, is anomalous in being recorded at only 50 m elevation in what is otherwise a montane species. It is in fruit only and thus remains doubtful taxonomically.

Evergreen shrubs or trees lacking a woody basal tuber, although occasionally thickened at base, mostly with inconspicuous lenticels on larger branches, lateral buds various, conspicuous to inconspicuous, scales rounded to acicular. Leaves entire or crenulate. Inflorescences of solitary racemes, mostly from leafy axils, few- to many-flowered. Flowers 5-merous with 10 stamens, anthers with tubules opening by introrse oblique or terminal pores which do not taper distally. Ovary 5-celled, but with thick intrusive parietal partial septa which give the appearance of a 10-locular ovary, each chamber containing several ovules. Fruit a spherical berry passing from green to blue or black when fully ripe.

The characters in Nakai's (1927) diagnosis have been greatly modified in the definition of this section used by Sleumer in his Vaccinioideen-Studien (Reference Sleumer1941). The bracts are described as often foliaceous and quite persistent in Nakai & Koidzumi (Reference Nakai and Koidzumi1927), whereas in the Flora Malesiana account (Reference Sleumer and Van Steenis1966–1967), they are described as “whether or not foliaceous or persistent”. Furthermore, Sleumer reduces sect. Euepigynium Schltr. with early deciduous bracts into synonymy in Flora Malesiana. Many of the Malesian species included in sect. Bracteata have bracts which vary from leaf-like and fully persistent to those which are minute, extremely reduced and very early caducous. In many cases, the bracts fall while the buds are very young and so have not always been described. Nakai's (1927) “biennial leaves” may be interpreted as perennial or evergreen which describes all the Bornean species. The lack of spurs on the anthers (Nakai, 1927) cannot be used to diagnose the section because the type species is described as often having two short spurs (Sleumer, Reference Sleumer and Van Steenis1966–1967). Similarly, the 10-celled ovary of Nakai (1927) is in fact a 5-locular ovary with five intrusive partial parietal septa (see Fig. 1A). Nakai does define the fruit colour as black (including blue here) which, as the ripe fruit colour, is considered a useful character in distinguishing this section from Rigiolepis in Borneo although red fruits do commonly occur in Vaccinium elsewhere. Vaccinium bracteatum in cultivation at the Royal Botanic Garden Edinburgh very occasionally produces two racemes from one leaf axil but this would appear to be a very rare occurrence in other species of Vaccinium sect. Bracteata. In Borneo, the anther tubules in this section do not significantly taper distally although in some other species in this section elsewhere (e.g. Vaccinium caudatum Warb.) they do. Vander Kloet & Dickinson (2009) have divided sect. Bracteata sensu Sleumer (Reference Sleumer and Van Steenis1966–1967) into several sections. No Bornean material was used in their analysis and the utility of this for identification purposes appears very limited. The species recorded here would occur at the end of their key (couplet 28) where only Vaccinium cercidifolium would fit their combination of characters of sect. Euepigynium Schltr. They place non-Bornean material of Vaccinium bancanum into sect. Eococcus Sleumer, but this species does not fit their key characters. A more detailed molecular analysis of the sections of Vaccinium is still awaited.

Chromosome number

2n = 24 (Atkinson et al., Reference Atkinson, Jong and Argent1995).

Distribution

Brunei. Indonesia: West Java, Sumatra, Anambas Islands, Banka, Billiton, Lingga, Kalimantan. Malaysia: Peninsula, Sabah and Sarawak.

Ecology

Common at sea level, possibly occurring to 1500 m. In sandy acid habitats, mostly terrestrial or occasionally epiphytic. Flowering: throughout the year but probably with seasonal flushes.

Conservation assessment

LC. A widespread and very common species.

Vaccinium bancanum Miq. and Vaccinium clementis Merr. appear to form a cline from lowland to montane habitats. Lowland forms with large leaves are easily placed in Vaccinium bancanum and montane forms with small leaves in V. clementis. The best distinguishing character appears to be the relative appearance of the resting lateral buds. In Vaccinium bancanum, they are mostly ovate to pyramidal, with broad scales, and are shorter than the subtending petioles (often only half the length). In Vaccinium clementis, they are very narrowly pyramidal with acicular, acutely pointed scales and are as long as or longer than the petioles that subtend them. This character may break down outside Borneo where material of Vaccinium bancanum var. kunstleri (King & Gamble) Sleumer from Peninsular Malaysia has been observed with narrowly pointed resting buds, but V. clementis is not known from this region and so is unlikely to cause confusion. Both species are very common and widespread in Borneo, and further observations are needed to see if this character works in all situations. Sleumer (Reference Sleumer and Van Steenis1966–1967, p. 796, couplet 97) uses confusing leaf shape characters and leaf length to distinguish the two species (more or less than 4 cm long). The leaf length will undoubtedly distinguish the lowland from the montane plants but does not help with intermediate specimens. Sleumer also gives a difference in rachis length in his descriptions: 3–10 cm in Vaccinium bancanum versus 1.5–4 cm in V. clementis. This nearly provides separation between these species but again probably represents the results of extremes of elevation. Further study is needed to see if these two species hybridise and whether any other characters can be used to separate them. There are some low-elevation specimens previously regarded as Vaccinium bancanum from kerangas (heath forest) with small leaves similar to those typical of V. clementis. These are now considered to be Vaccinium stenanthum Sleumer. A specimen from Mount Berumput in Sarawak, Argent 19942998, is unusual in having double spurs on the anthers, two pointing up and two pointing down. The glands on the anthers and the simple hairs inside the corolla are often very difficult to see; they lack contrast and the glands in some cases disappear with age. A specimen at Kew (Teijsmann s.n., K000780712), is cited as a doubtful isolectotype above, following Sleumer's annotation, “possibly part of the type coll.”

Vaccinium bancanum var. tenuinervium differs chiefly from the nominate variety in that the calyx lobes have several sessile, subpersistent lateral glands, and by the generally narrower, elliptic or oblong-lanceolate, more distinctly acuminate or subcaudate, often thinner leaves.

Distribution

Indonesia: Sumatra, West Java, Central Kalimantan. Malaysia: Peninsula and Sarawak.

Ecology

As for Vaccinium bancanum var. bancanum, Sleumer (Reference Sleumer and Van Steenis1966–1967) records this growing possibly as high as 2050 m. It is doubtful if it really occurs as high as this, because many specimens have been misidentified in the past. Flowering: January to December.

Conservation assessment

LC. A very common and widespread variety.

2. Vaccinium ceraceum Argent, sp. nov.

Vaccinium ceraceum is distinct in the following combination of characters: having the inflorescence covered with tufts of filamentous wax which appear as grey scurfy patches when dry, and the persistent bracts having prominent sessile glands on the margins. Superficially similar to Vaccinium simulans but the leaves are smaller, with the venation more pronounced abaxially rather than adaxially. Also differing from Vaccinium simulans and other similar species in having only minute patent hairs on the filaments, unlike the much longer semi-appressed hairs on the filaments of those species, and a style which is glabrous except for a papillose area near the apex. – Type: Indonesia, East Kalimantan, Krayan, Long Bawan, en route from Long Bawan to Long Umung, 15 ix 1990, M. Kato, H. Okada, J. Murata, D. Darnaedi, H. Akiyama, Y. Watano & D. Komara 23426 (holo BO; iso E, L, TI). Figs 31, 32.

Fig. 31.

Vaccinium ceraceum Argent. Kato et al. 23426. A, Habit (scale bar, 3 cm); B, flower (scale bar, 2 mm); C, bract and bracteole (scale bar, 2 mm), with inset showing bract margin (scale bar, 0.5 mm); D, stamens (scale bar, 1 mm), with inset showing hairs on filament (scale bar, 0.2 mm); E, pistil with calyx (scale bar, 1 mm). (Illustration: Claire Banks.)

Fig. 32.

Vaccinium ceraceum. Type specimen, Kato et al. 23426. A, Leaf surface, showing tufts of wax; B, bract margin, showing large glands. (Scanning electron micrographs: Frieda Christie.)

Terrestrial small tree, c.4 m. Twigs minutely patently pubescent, quickly glabrescent, leaves subdensely arranged; lateral buds c.2 × 1 mm, broadly conical moderately conspicuous, almost as long as subtending petioles, scales subtriangular caudate, hairy especially on margins. Leaves: petiole c.2 × 1 mm, minutely hairy, flattened; blades 25–45 × 14–18 mm, elliptic, base tapering and narrowly decurrent into petiole, margin entire, narrowly cartilaginous, flat or weakly revolute, with 1 or 2 marginal glands on each side, 1–3 mm from petiole, lamina with brown glandular hair bases beneath, apex shortly (3–5 mm) apiculate, extreme tip obtuse; midvein narrowly and weakly impressed above, very weakly raised below; lateral veins 2–4 each side, 2 or 3 high-arching from near base, often with one from well above base, faint, reticulation inconspicuous. Inflorescence of solitary racemes, in upper axils, 6- to 8-flowered, flowers secund, rachis c.20 mm, glabrous proximally but with lax covering of patches of grey, scurfy indumentum distally, bracts to 35 × 1.25 mm, elliptic, often with short acuminate scaly point, mostly persisting to anthesis, with distinct globose glands, unlike the more usual club-shaped glands where these occur in other species, and some short simple hairs on margins; bracteoles basal, minute, subulate, persisting to anthesis, fringed with simple white hairs. Flowers: pedicels 2–3 × c.0.4 mm, scaly with pale grey wax; calyx densely covered with tufts of filamentous wax, tube campanulate, 0.75 × 1.25 mm, lobes triangular, c.0.5 × 1 mm, fringed with some short simple hairs but lacking both apical and marginal glands; corolla c.4 × 2.5 mm, barrel-shaped, pink, glabrous outside but covered with tufts of filamentous wax, laxly hairy proximally inside, lobes c.0.7 mm, reflexed; stamens: filaments 2 mm, glabrous, anther cells c.0.5 mm, with short awns, tubules c.0.2 mm, at right angles to cells, with small glandular hairs when young, pores circular; disc glabrous; style c.2 mm, cylindrical, glabrous but papillose near apex. Fruit not seen.

Distribution

Indonesia: East Kalimantan. Known only from the type collection.

Ecology

Collected at 900 m, but the vegetation type not recorded. Flowering: September.

Etymology

From the Latin – ceraceus – waxy. An allusion to the distinctive waxy patches on the inflorescences of this species.

This is a curious and very distinctive species with tufts of filamentous wax on the inflorescences which appear to the naked eye as a grey scurfy indumentum. Filamentous wax has been recorded elsewhere on the fruits of Vaccinium (Chu et al., Reference Chu, Gao, Cao, Fang, Chen and Xiao2017) but does not appear to have been recorded from any other of the Southeast Asian species. This feature disappears on boiling with detergent and will also probably disappear when collections are made into alcohol. A more permanent feature is the appearance of the bracts, which have large sessile glands on the margins similar to those often described from the margins of the calyx lobes of several other Southeast Asian Vaccinium species.

This new species looks superficially similar to Vaccinium simulans, but the leaves are smaller with the venation more pronounced abaxially rather than adaxially (the opposite of the situation in V. simulans). It differs also from Vaccinium simulans and most other similar species in having minute hairs that are very difficult to see on the filaments rather than being glabrous or with distinct long hairs. In addition, apart from papillae near the stigma, the style is glabrous. Vaccinium elliptifolium differs in from V. ceraceum in its prostrate shrubby habit (versus tree) and having more distinct lateral veins and longer pedicels.

Fig. 33.

Vaccinium cercidifolium. RBGE accession number 19820845; Gunung Bungo Range, Sarawak. (Photograph: G. Argent.)

Terrestrial or occasionally epiphytic shrub to 2 m. Twigs glabrous, or (? see comment below) patently short pubescent, rounded, green often flushed with red, leaves arranged in lax spirals, lateral buds up to 7 × 2 mm, conspicuous, narrowly conical, as long as or slightly longer than petioles. Leaves: petiole up to 5 × 3 mm, strongly flushed red when fresh, rugulose, not or only weakly grooved above, initially covered with brown glandular hairs, blade 35–85 × 25–55 mm, broadly elliptic to subcircular; base rounded to cordate, margin entire, flat or narrowly revolute when dry, narrowly cartilaginous, with 2 marginal glands c.1 mm from petiole, apex very broadly pointed to rounded, minutely brown glandular hairy on both sides when young, quickly glabrescent, midvein adaxially, broadly raised for up to 10 mm, red and triangular for up to 10 mm, then impressed distally for remainder of length of leaf, abaxially broadly raised for most of length of leaf, lateral veins 1–3 pairs, basal pair short, disappearing within 10–20 mm, second pair arising close to base, strong, curving upwards and disappearing in upper part of leaf, a further pair or pairs arising from basal half of leaf, curving upwards and disappearing before margin: reticulation obscure in fresh leaves, prominent on both sides when dry. Inflorescence of racemes from upper axils, with 10–20 secund flowers; rachis 80–100 mm, glabrous; bracts up to 20 × 8 mm, elliptic to elliptic-obovate, distinctly apiculate, entire, glabrous, pale green, falling very early, often even before inflorescence has elongated; bracteoles minute, up to 1 mm, filamentous, glabrous, withering and disappearing as bracts fall. Flowers: pedicels 10–12 × c.1 mm, glabrous; calyx hemispherical, glabrous, tube to 2 mm long, limb slightly spreading to c.1 mm, lobed nearly to base, lobes ovate-triangular, obtuse, lacking or with only a minute terminal gland, without marginal glands; corolla 14 × 9 mm, ellipsoid, red, pale pink, or pale brownish cream; scented with heavy honey scent; lobes c.2.5 × 1.2 mm, broadly triangular, minutely papillose and with a few long white hairs, reflexed; stamens: filaments from a broad densely hairy base (c.0.9 mm wide) becoming very slender distally (c.0.1 mm) and less hairy, c.5 mm long; anthers with tubules bent at right angles to cells; cells 1.2 mm, with a pair of distinct awns at apex, tubules with several conspicuous glands 0.9 mm, widely diverging, pores truncate, circular; disc laxly hairy; style 10–12 mm, with dense appressed hairs in proximal 3/4, glabrous distally. Fruit not recorded.

Distribution

Indonesia: Central Kalimantan. Malaysia: Sarawak, Mount Dulit, Mount Buri. Widespread and probably under-recorded.

Ecology

Lower montane forest, often on ridges at 600–1700 m. Flowering: March, September to December.

A distinctive species with prominent fan-like leaf venation and rounded to cordate leaf bases which are dark red when fresh and dry dark brown. The two original collections were said to have “blood red flowers”. The specimen cultivated in Edinburgh (RBGE accession number 19820845) from the Gunung Bungo Range, Sarawak has creamy brown flowers (see Fig. 33). Sleumer (Reference Sleumer and Van Steenis1966–1967) describes the twigs as “patently short pubescent”, while the cultivated material in Edinburgh and most of the specimens examined have glabrous twigs.

Fig. 34.

Vaccinium claoxylon. RBGE accession number 19801232; Mount Kinabalu, Sabah. (Photograph: G. Argent.)

Terrestrial shrub or occasionally small tree to 10 m, rarely epiphytic. Twigs glabrous, red when young with prominent pale lenticels, leaves in clusters separated by bare stems from which deciduous scale leaves have fallen, older stems with conspicuous, distinctly raised, subcircular leaf scars, lateral buds very prominent, hemispherical, up to 4 × 4 mm with tightly appressed rounded scales. Leaves: petiole very short, almost obsolete on Kinabalu, 0–1 × 4–5 mm (up to 6 mm long in East Kalimantan), bright red; blade 70–140 × 40–60 mm, elliptic, broadly elliptic to obovate, base narrowly tapering, often decurrent, margin entire, narrowly cartilaginous, flat or weakly revolute, with a marginal gland on each side at base which protrudes beyond margin, lamina glabrous above, laxly glandular hairy beneath when young, apex obtuse, rounded, very shortly and broadly acuminate or rarely weakly retuse, midvein red near base, raised in proximal half above, then impressed distally, raised throughout beneath, as broad as petiole at base, tapering gradually upwards; lateral veins 4–8 per side, pinnate, emerging from midvein at an acute angle, curving distally and disappearing before the margin, reticulation dense and prominent when dry. Inflorescence of racemes, mostly emerging below leaves from defoliate axils, stiffly semi-erect, flowers secund, rachis 60–130 mm, glabrous, bracts c.6 × 2 mm, narrowly ovate, with a few irregular marginal glands, minutely punctate abaxially, deciduous well before flowers open; bracteoles 1 × 0.5 mm, with prominent sessile glands all round margin, inserted just below calyx. Flowers: pedicels 1–2 mm long, thick, glabrous: calyx tube hemispherical 1–1.5 mm, limb erect, lobes ovate-triangular 1.2–1.5 mm, with numerous small thick, sessile glands and some fine ciliate hairs along margin; corolla 6–8 × 2.5 mm, narrowly tubular-urceolate, glabrous on both sides or hairy proximally within, pale to dark pink, often paler proximally, strongly almond scented, lobes ovate, becoming reflexed, 1 mm, pale pink to almost white; stamens c.4 mm, becoming slightly exserted after corolla lobes reflex, slightly unequal; filaments dilated just above base, becoming slender distally, long-hairy in proximal 1/3 to 2/3, glabrous distally; anther cells c.1.3 mm including very short tubules, diverging from base, oblongoid, slightly granular, without spurs; tubules as wide as cells, with oblique pores; disc prominent, glabrous, style 7–10 mm, columnar, broadened at base, densely covered with long distally pointing appressed hairs in proximal 2/3, glabrous distally, becoming exserted from corolla. Fruit spherical, green often with a red or purplish flush when immature, ripening black.

Distribution

Brunei. Indonesia: Kalimantan. Malaysia: Sabah and Sarawak. Widespread on mountains in Borneo.

Ecology

Submontane mossy forest and shrubberies 700–3000 m. Flowering: January to September.

Conservation assessment

LC. Common on many mountains in Borneo, several of which have protected status.

This species is distinct with its prominent, pinnate, translucent venation, twigs with very conspicuous lenticels (unusual in this section) and raised leaf scars. The inflorescences also arise from the axils of fallen scale leaves, thus mostly from bare stems below or between the foliage leaves, which is also unusual for species in sect. Bracteata. It is unlikely to be mistaken for a member of Rigiolepis, however, because of its rigid, erect, usually terrestrial habit and relatively large flowers. The extremely divergent anther cells are quite unlike those of any other Bornean species.

Fig. 35.

Vaccinium clementis. Cultivated, RBGE accession number 19801294; Mount Kinabalu, Sabah. (Photograph: G. Argent.)

Terrestrial shrub or small tree up to 12 m, occasionally epiphytic. Twigs minutely patently pubescent, or glabrous, with occasional brown glandular hairs, leaves subdensely arranged, lateral buds 1.5–3 mm, very narrowly conical, conspicuous to obscure, mostly as long as or longer than subtending petioles, scales subulate. Leaves: petiole 1–2 × 1–1.5 mm, minutely hairy, without a groove, blades 20–30 × 7–15 mm, elliptic to weakly obovate, base tapering, with 1 or 2 marginal glands on each side, 2–3 mm from petiole, margin entire, narrowly cartilaginous, flat or weakly revolute, lamina with brown glandular hairs beneath when young, apex rounded to obtuse, sometimes broadly contracted apically, midvein smooth to weakly impressed above, smooth to very weakly raised below; lateral veins 1 or 2, each side high-arching from near base, faint, reticulation inconspicuous. Inflorescence of solitary racemes, in upper axils, with 6–12 secund flowers; rachis 20–50 mm, minutely hairy, bracts up to 10 × 5 mm, curling back and mostly falling well before flowers open; bracteoles minute, subulate, withering and falling early. Flowers: pedicels 5–8 mm, minutely hairy; calyx glabrous, or shortly hairy, tube subglobose, base 2.5–3 mm in diameter, lobes triangular, c.1 mm, with an apical gland and some marginal sessile glands; corolla 7–8 × 5–6 mm, barrel-shaped or weakly conical, white, through pink to red, faintly scented, glabrous outside, laxly hairy proximally inside, lobes c.1 mm, reflexed; stamens 3–4 mm, filaments 2.5–3 mm, broad at base, narrowing upwards to become very slender, densely hairy in proximal 2/3; anthers 0.8–1 mm, cells 0.6–0.8 mm, mostly with 2 short awns, tubules c.0.2 mm, with small glandular hairs when young, these disappearing often by anthesis, pores circular, opening slightly obliquely; disc densely hairy; style 7–8 mm, densely hairy in proximal 3/4. Fruit c.5 mm in diameter, spherical, shiny black when ripe.

Distribution

Brunei. Indonesia: Kalimantan, less common in the east. Malaysia: widespread in both Sabah and Sarawak.

Ecology

Mostly terrestrial in submontane and montane mossy forest, 600–2400 m. Flowering: January to December but with seasonal flushes.

Conservation assessment

LC. Widespread and common, its montane habitat being less threatened at present from disturbance.

Vaccinium clementis is a common montane species which is very variable owing to the broad range of habitats which it can occupy. At the upper end of its elevational range, it has been confused with Vaccinium coriaceum. This species usually has smaller less obovate leaves than Vaccinium clementis but there is little florally to separate these two species. Occasionally, the racemes terminate in a peloric flower with up to double the usual number of parts.

At the lower end of its range, it can be difficult to tell from Vaccinium bancanum (see under that species), but V. bancanum usually has more acuminately attenuate leaf apices and conical lateral buds which are much shorter than the petioles, at least in Borneo. Vaccinium elliptifolium Merr. can also be difficult to separate from V. clementis (see notes under Vaccinium elliptifolium). Clemens 51648 from the Mesilau River in Kinabalu is a poor specimen without flowers and apparently not seen by Sleumer. Although the leaves do not have any marginal glands (contrary to the information on the label), the size and shape of the leaves agree with Vaccinium clementis.

Fig. 36.

Vaccinium cordifolium. Mount Kinabalu, Sabah. (Photograph: G. Argent.)

Shrub 1–2 m, branches arching and often becoming pendent. Twigs densely patently grey or brown hairy when young becoming glabrescent; leaves crowded, dense and overlapping at tips of branches, lateral buds 2.5–3 × 0.5–0.8 mm, narrowly conical, scales acicular. Leaves: petiole 2–3 × 1.5–2 mm, hairy, blade 24–45 × 15–30 mm, ovate, subsessile, base auriculate, often slightly amplexicaul, margin flat, entire, or very slightly revolute, marginal glands small, a single pair on the auricles, c.2 mm from petiole, apex bluntly attenuate, dark green and glossy above, paler beneath, patently shortly pubescent on both sides when young, quickly glabrescent except on basal part of lamina and midvein beneath, midvein level above, prominent beneath, lateral veins 6 or 7 per side, 2 or 3 basal, distal ones from midvein, curved, ascending towards margin, slightly raised above, more conspicuously so below, reticulation dense, finely prominent beneath. Inflorescence of racemes from uppermost axils, solitary or rarely in pairs, laxly 5- to 12-flowered, flowers secund; rachis 20–60 mm, densely white pubescent, bracts up to 8 × 2 mm, ovate, leafy, falling before anthesis; bracteoles not seen. Flowers: pedicels 8–12 mm, slender, densely white pubescent: calyx 1–1.5 mm, reddish or purplish, tube widely campanulate, tomentose, limb erectopatent, 5-lobed nearly to base, lobes ovate-acuminate, with conspicuous subparallel dorsal veins, laxly pubescent and with a ciliate margin, c.3 × 2 mm, without distinct apical gland; corolla 12–15 × 5–6 mm, tubular-urceolate, white usually strongly flushed with pink, or pink with white lobes, glabrous on both sides, lobes 1.5 mm, ovate, obtuse, reflexing; stamens subequal, often incompletely developed, filaments 6–7 mm, linear-subulate, slightly dilated just above base, densely long hairy except just above base, anther cells oblongoid, c.2 mm, with 2 very short dorsal spurs, tubules cylindrical, much narrower than cells, c.1.5 mm, divergent, laxly covered with short gland-tipped hairs, pore subcircular, flaring; disc densely covered with long erect hairs; style red, columnar, densely subappressed hairy in lower 2/3, 10–11 mm. Fruit up to c.12 mm in diameter, a globose berry, greenish, often flushed with red on one side, ripening purplish-black with persistent, leathery calyx lobes.

Distribution

Malaysia: Sabah, Mount Kinabalu and Mount Trus Madi. Common on both mountains but yet to be recorded elsewhere in Borneo.

Ecology

In dwarf subalpine forest and submontane shrubberies on open ridges, 2400–2900 m, on peat overlying granite and ultramafic rocks. Flowering: January to December.

Conservation assessment

LC. Both known localities have good protection. It is a common species along the main trail up Mount Kinabalu, occurring in abundance.

A very distinctive species with its conspicuously white hairy stems and strongly auriculate leaf bases.

Chromosome number

2n = 24 (Atkinson et al., Reference Atkinson, Jong and Argent1995).

Distribution

Malaysia: Sabah, Mount Kinabalu.

Ecology

Montane forest and shrubberies and open granite rocky areas, 2000–3400 m. Flowering: January to December.

Conservation assessment

LC. Substantial populations in Gunung Kinabalu National Park.

See discussion following the account of var. stapfianum.

Differing in the hairy calyx from the type variety.

Distribution

Indonesia: East (?) Kalimantan. Malaysia: Sarawak. First described from Gunung Mulu in Sarawak and subsequently found to be widespread on mountains in Borneo.

Ecology

Montane forest, 1500–2500 m elevation. Flowering: January to December.

Conservation assessment

LC. Its widespread distribution and occurrence in several national parks would indicate that there are abundant, well-protected populations.

Distinctly epiphytic, with laxer leaf disposition than the type and with shorter, smaller flowers, 2.5–3 mm long.

Distribution

Malaysia: Sabah, Mount Kinabalu only.

Ecology

Epiphytic in mossy montane forest, 1500–2700 m. Flowering: February to July, probably during other months.

Conservation assessment

LC. Although recorded only from Gunung Kinabalu, it is widespread on this mountain, which is well-protected.

The taxonomic value of recognising var. minus is doubtful because, as has already been noted, the flowers of this species are very variable in size. The differences in the leaves and flower size are no doubt due to less exposure in the shady epiphytic habitat. It would be interesting to see if the differences were maintained in cultivation experiments.

Fig. 37.

Vaccinium coriaceum var. stapfianum (Sleumer) Argent. RBGE accession number 19801165; Mount Kinabalu, Sabah. (Photograph: G. Argent.)

Differing from the previous varieties in having an entire leaf margin with only a single pair of impressed glands (more rarely 2 or 3 pairs of glands) in the proximal half of the leaf and a glabrous or only very laxly hairy calyx.

Distribution

Malaysia: Sabah, Mount Kinabalu and the Crocker Range.

Ecology

Terrestrial on both ultramafic and granite areas of Gunung Kinabalu, 2000–3600 m elevation. Flowering: throughout the year.

Conservation assessment

LC. An abundant species in Gunung Kinabalu National Park, where it is protected.

Vaccinium stapfianum Sleumer is here recognised as a variety within Vaccinium coriaceum Hook.f. (which has four varieties in Borneo). There is great variation in the number and distribution of marginal leaf glands in Vaccinium coriaceum although these may be constant within local populations. In the absence of other differentiating characters, Vaccinium stapfianum would appear better at varietal rank. Vaccinium coriaceum var. hirsuticalyx Argent, with a densely hairy calyx, is the common variety away from Kinabalu, mostly having just one pair of glands on the leaf margin, slightly proximal to the middle of the leaf. The specimen SAN.141808 from Long Miau, Meligan, Sipitang District, Sabah has the densely hairy calyx but the corolla is glabrous inside.

Vaccinium coriaceum is very similar to the widespread Vaccinium lucidum (Blume) Miq., not yet recorded in Borneo. Vaccinium coriaceum is the older name if these species were to be united. Vaccinium timorense Fawc., with similar small leaves, is widespread further to the east. It is a curious anomaly that Sleumer (Reference Sleumer1966–1967) included Vaccinium timorense var. denticulatum Fawc. [subcrenulate with glands all round the leaf margin] within V. timorense [with a single pair of basal glands], not recognising it at even varietal rank, whereas V. coriaceum and V. stapfianum were separated on just this character. Other records of what are here considered to be Vaccinium coriaceum have been collected from widespread localities in Borneo. Most of these have been previously identified as Vaccinium clementis. The variation accorded here to the presence and number of the marginal glands in Vaccinium stapfianum casts some doubt on the validity of Vaccinium leptocladum Sleumer as a good species. See discussion under that species.

Fig. 38.

Vaccinium costerifolium. Argent et al. 9661; Nyaru Menteng, Kalimantan. (Photograph: G. Argent.)

Trailing or scrambling shrub with arching stems to 2 m. Twigs rounded, sparsely covered with lenticels, minutely patently hairy, laxly leaved, lateral buds to 2 mm, ovoid-conical with acute apex, inconspicuous. Leaves: petiole 2–3 mm, slender, rugulose, minutely hairy, blade 25–60 × 12–30 mm, elliptic, base broadly attenuate into slender petiole, margin entire, slightly revolute, distinctly so when dry, with 1 or 2 small marginal glands, a little above base on each side, sometimes with a few additional upper ones, glabrous minutely laxly glandular hairy underneath, apex shortly gradually and obtusely attenuate, midvein impressed above, prominent beneath, lateral veins 2 pairs, curving upwards from or near the base, 1 or 2 upper shorter curving veins, all impressed above, prominent beneath, reticulation slightly raised on both sides or obscure. Inflorescence of racemes from axils in younger part of twigs, 8- to 12-flowered, flowers secund, rachis slender, angular, minutely patently hairy, bracts sometimes persisting until anthesis, bracts ovate, 3–6 × c.2 mm, with distinct veins, fringed with minute hairs and punctate underneath; bracteoles subulate, basal, to 1.8 mm, minutely hairy. Flowers: pedicels slender, 3–6 mm, minutely and sparsely patently hairy; calyx tube rounded, 1–1.5 mm, shortly densely hairy and laxly glandular hairy, limb suberect, deeply divided, lobes triangular, 2–2.5 mm, less densely hairy than calyx tube, without apical gland: corolla 8–10 × c.4 mm, tubular, slightly contracted distally, with distinct angles in bud, membranous, pink to red to deep purple, with white tips to the lobes, sweetly scented, minutely hairy outside with both simple and glandular hairs and at base inside, lobes c.1 mm, obtuse, suberect; stamens: filaments c.3 mm, base broad, hairy, tapering rapidly to slender apical part which is glabrous distally; anther cells 1.2–1.3 mm, oblongoid, granular, shortly obtusely 2-spurred dorsally; tubules cylindrical, narrower than cells, very slightly dilated upwards, held at right angles to cells, c.1.4 mm, subobliquely opening apically, back of each tubule extended into 1 or 2 short recurved teeth; disc prominent, glabrous or laxly hairy; style 9–10 mm, columnar, glabrous becoming exserted as flower ages. Fruit subglobose, c.6 mm in diameter, laxly hairy, green, flushed red, turning black.

Distribution

Indonesia: South and Central Kalimantan.

Ecology

Near sea level in swampy or open heath forest on acid sandy or peaty soil. Locally common. Flowering: January and October. Fruiting: September.

Conservation assessment

LC. Although this species is not known from many localities, the recent collection (Argent et al. 9661) from Nyaru Menteng was in poor swampy ground unlikely to be developed for agriculture and was collected from a substantial population. Kessler 1587, however, was growing among pineapples in a cultivated area.

The original description is here amplified from Argent et al. 9661.

Erect shrub or small tree. Twigs rounded, densely covered with subpatent hairs, densely to laxly leaved, lateral buds obscure to prominent, up to 4 mm, slender, scales subulate, hairy abaxially. Leaves: petiole 2–3 × 1–1.8 mm, densely hairy; blade 20–32 × 12–24 mm, elliptic to broadly elliptic, base broadly attenuate to rounded, margin subcartilaginous, entire, narrowly revolute, marginal glands a single pair at 2–5 mm from petiole, hairy at base and lower half of midvein below and laxly glandular punctate, apex retuse, sometimes rounded, then usually with small central cleft which may have an indentation either side of it (? glands), midvein impressed above and prominent below, lateral veins mostly 2 pairs at an acute angle; one pair from base ascending into upper part of leaf but disappearing before margin, upper pair shorter but similar, reticulation mostly obscure. Inflorescence of racemes from upper axils, flowers 6–10, secund; rachis 20–25 mm, rounded, densely white-hairy; bracts 3–4 mm, narrowly cordate, glabrous except for a fringe of white hairs; bracteoles 1.5–2 mm, subulate, basal, subpersistent, white-hairy especially towards the base. Flowers: pedicels 3–5 mm, densely white-hairy; calyx tube globose, densely hairy, limb erect, deeply lobed, lobes triangular, ovate to acute, glabrous or very sparsely hairy abaxially, with a dense fringe of white hairs on margin, mostly without conspicuous marginal or terminal glands; corolla urceolate 7–9 × 3 mm, glabrous outside, long-hairy inside, red; lobes c.1 mm, ovate, recurved with minutely denticulate margin; stamens: filaments 1.5 mm, subulate, long-hairy; anther cells 0.5 mm, oblongoid, without spurs, tubules up to 0.3 mm, as wide as cells, truncate, pore round margin reflexing and with a few gland-tipped hairs; disc densely hairy; style 8–9 mm long-hairy in proximal 1/2 to 2/3. Fruit (immature) globose, remaining densely hairy and with persistent erect calyx lobes.

Distribution

Malaysia: Sabah, Mount Kinabalu.

Ecology

In submontane forest in ultramafic areas, 1200–2300 m elevation. Flowering: May to July.

Conservation assessment

LC. Although known only from a single mountain, there are good populations of this species in several different localities on Gunung Kinabalu, where it grows in well-protected forest sites. It is common alongside the recently developed alternative trail up the mountain from the Mesilau rest house.

This species is superficially similar to Vaccinium clementis but the hairs on the stems and inflorescences are much more obvious, being longer and coarser than those on the stems of V. clementis, which has glabrous or very finely and shortly tomentose stems and inflorescences. The leaf apices are characteristically rounded with a retuse tip and sometimes with two characteristic depressions on either side of the apex, whereas Vaccinium clementis has leaves that are mostly broadly acuminate with a rounded but not retuse apex. The calyx lobes on the fruit are also different, being erect in Vaccinium elliptifolium and normally appressed to the disc in V. clementis. Material of Vaccinium elliptifolium usually dries darker than specimens of V. clementis, and V. elliptifolium would appear to be restricted to ultramafic habitats on Mount Kinabalu and its environs whereas V. clementis grows in the more usual acid habitats of this family.

The two Clemens collections cited by Merrill in the protologue of this species, 10894 and 11099, are presumed to have been destroyed in Manila in the Second World War. Sleumer (Reference Sleumer1961) saw neither collection but still selected Clemens 10894 (PNH) as the lectotype. No duplicates of these collections, which are the original material, have been located. Three other Clemens collections were made in the type locality, namely 31063, 31404 and 40194. None of these has been seen but duplicates may well turn up in other herbaria and would then be candidates for neotypification.

Non Vaccinium endertii (J.J.Sm.) Masam. = Rigiolepis endertii J.J.Sm.

Shrub to 6 m, mostly epiphytic. Twigs thick, at first angular, glabrous, leaves lax, lateral buds 5–6 mm, ovoid-oblongoid, scales ovate, acute. Leaves: petiole 4–5 × c.1.5 mm, glabrous, blade 45–95 × 25–43 mm, obovate to elliptic, base rounded to subcordate, marginal glands at a short distance from petiole, glabrous except for lax glandular hairs below which disappear leaving surface punctate, margin entire, apex broadly attenuate, obtuse to rounded, midvein impressed above, strongly prominent beneath, lateral veins 3 or 4 from base and slightly above base, curving upwards and with a few shorter ones from distal part of leaf, reticulation lax, inconspicuous, slightly raised beneath. Inflorescence of 8- to 15-flowered racemes with laxly secund flowers; rachis 50–83 mm, angular, glabrous; bracts and bracteoles not recorded. Flowers: pedicels 9–12 mm, glabrous; calyx tube cup-shaped, rugulose, 2 mm in diameter, limb spreading, deeply lobed, lobes 1.5 mm, subtriangular, sometimes obscurely ciliate, apex obtuse with a thick gland; corolla 10–12 × 3–4 mm, tubular-urceolate, greenish white with red base, glabrous outside, subdensely hairy in proximal half inside; lobes 1.5 mm, obtuse, becoming reflexed; stamens: filaments 3–4 mm, linear with base dilated, long hairy; anther cells 1.5 mm, oblongoid, echinulate, distinctly 2-spurred dorsally; tubules c.1 mm, with a few gland-tipped hairs, cylindrical, narrower than cells and at a wide angle to them, opening transversely; disc hairy; style 9–10 mm, columnar, densely hairy with simple white hairs in basal 3/4. Fruit globose, c.8 mm in diameter (dry).

Distribution

Indonesia: Central Kalimantan, Amai Ambit and Mount Kemel. Malaysia: Sarawak, Upper Mujong area.

Ecology

In primary ridge forest, at c.1000 m. Flowering: October.

This is a poorly known species that requires further collection.

Slender epiphytic shrub to 1.5 m. Twigs slender, up to 2 mm in diameter, angular, patently very shortly hairy, laxly to subdensely leafy, lateral buds inconspicuous. Leaves: petiole c.1 mm long, slender; blade 7–10 × 2.5–3 mm, narrowly obovate to elliptic, base tapering, margin entire, revolute, with only an occasional marginal gland c.1 mm from pedicel, lamina with relatively large, dark gland bases abaxially; apex rounded; midvein slightly raised on both side, lateral veins 2 or 3 arising from base or slightly above base, ascending to upper part of leaf, slightly raised on both sides, reticulation dense, only slightly raised on both sides. Inflorescence of racemes from upper axils, 1- or 2-flowered; rachis 2–5 mm long, glabrous; bracts and bracteoles not recorded. Flowers: pedicels 5–7 mm, glabrous; calyx tube 0.5 mm, broadly campanulate, densely pubescent, limb suberect, deeply 5-lobed, lobes 1 mm, triangular with obtuse apex, margins ciliate and with several thick, subsessile glands; corolla 3.5–4 × c.2.5 mm, red, ovoid-urceolate, inflated below, abruptly contracted distally, glabrous outside, shortly hairy towards base inside; lobes up to 1 mm; stamens: filaments 1.5 mm, linear, long-hairy; anthers c.0.8 mm, cells broadly oblongoid, without spurs, tubules very short, as wide as cells, opening by oblique pores; disc laxly hairy; style 4 mm, filiform, hairy in proximal half. Fruit not seen.

Distribution

Indonesia: East, Kalimantan, West Kutei, Mount Kemul.

Ecology

Montane forest, 1600 m. Flowering: October.

This species is known only from the type collection. It is very similar to Vaccinium coriaceum but described with a much shorter inflorescence which is only 1- or 2-flowered. Contrary to the original description, there are occasional marginal glands on a few leaves, although on most they are completely lacking. Further collections of this species from the type locality are very desirable.

Epiphytic climbing shrub. Twigs angular, densely shortly pubescent when young, subdensely spirally leaved, lateral buds subglobular, 2–3 mm in diameter, scales broadly ovate and obtuse. Leaves: petiole 3–4 × 1–1.5 mm, somewhat flattened, grooved above, shortly hairy when young, blade 50–72 × 20–32 mm, elliptic or obovate-elliptic, base tapering, apex obtuse to rounded, margin entire, somewhat revolute, the pair of marginal glands obscure, shortly pubescent and laxly glandular hairy when young beneath, midvein narrowly impressed especially towards base above, strongly obtusely prominent beneath, lateral veins 1 or 2 basal and 5 or 6 upper pinnate, suberect to widely spreading pairs, a little raised on both sides, reticulation lax and faintly visible. Inflorescence of 15- to 25-flowered racemes from upper axils; rachis 30–35 mm, slender, densely shortly pubescent; bracts up to 5 × 4 mm, subcircular to ovate; bracteoles not seen. Flowers: pedicels c.1.5 mm, curved, densely shortly pubescent: calyx tube c.1 mm, campanulate, laxly hairy at base, limb spreading, c.1 mm, lobes triangular, with thick apical gland; corolla c.4–5 × 2 mm, thin, glabrous outside, laxly hairy in lower half inside, lobes recurved, very short; filaments 1.8 mm, from a broad base becoming linear, subdensely hairy; anther cells 0.5 mm, oblongoid, with 2 short dorsal spurs; tubules cylindrical, as wide as and as long or a little longer than cells, with a few short gland-tipped hairs, opening transversely and sometimes with back wall produced into a minute tooth; disc hairy; style c.2.5 mm, hairy in proximal half. Fruit not seen.

Distribution

Indonesia: Central Kalimantan, Mount Batu Tiban, East Mount Kemel.

Ecology

Epiphytic, climbing shrub in primary forest, 1700–1850 m.

This species is apparently known from only two collections. Endert s.n. (SING), from the type locality, was collected soon after the type collection and is similar. The flower colour is not recorded, and the indication of a climbing habit may be doubtful because epiphytes are often mistaken for climbers.

Fig. 39.

Vaccinium pachydermum. Gunung Mulu, Sarawak. (Photograph: G. Argent.)

Erect shrub or small tree, up to 6 m. Twigs thick, shortly densely hairy becoming glabrescent, laxly to subdensely leaved; lateral buds ovoid to spherical, obtuse, scales ovate, up to 2 mm. Leaves: petiole 5–10 × 2–4 mm, densely pubescent; blade 35–100 × 25–70 mm, elliptic, broadly elliptic, obovate-elliptic to subcircular, base tapering and decurrent, margin entire, recurved when dry especially towards the base, usually with a pair of marginal glands 1–1.5 mm from petiole, glabrous above, laxly glandular hairy beneath when young, abaxial glands leaving a punctate surface when gone; apex mostly rounded or broadly obtuse, occasionally broadly acuminate; midvein impressed above, very prominent beneath, lateral veins 2 or 3 basal and somewhat above base, and additionally 2 or 3 upper pairs all ascending or with all lateral veins inconspicuous, reticulation obscure. Inflorescence of lax many-flowered racemes from upper leafy axils, flowers secund; rachis 40–80 mm, densely hairy; bracts up to 8 × 3 mm, narrowly cordate to elliptic, glabrous, falling early; bracteoles minute, slightly suprabasal, subulate. Flowers: pedicels 6–12 mm (to 20 mm in fruit), densely hairy, curved; calyx tube hemispherical, 2 mm, limb spreading, 2–2.5 mm, 5-lobed almost to base; lobes ovate-triangular, less pubescent than calyx tube, ciliate, with a small apical gland and often several sessile marginal glands; corolla 8–9 × 4–6 mm, broadly tubular-urceolate, pink or white, sweetly scented, glabrous to laxly hairy outside, laxly hairy at base and near opening inside; lobes 1.5 mm, recurved, often flushed pink; stamens: filaments c.3 mm, linear, densely hairy; anther cells 1.5 mm, oblongoid, echinulate with 2 dorsal spurs; tubules c.1 mm, cylindrical, as wide as cells, erect, opening transversely and with several gland-tipped hairs; disc densely hairy; style c.8 mm, slender, hairy in proximal 3/4. Fruit 6–8 mm in diameter, globose with a flat top, ripening blue, remaining pubescent with erect calyx lobes and a prominent disc.

Distribution

Brunei: Mount Pagon. Malaysia: Sabah, Mount Kinabalu; Sarawak, Mount Kalulong, Mount Dulit, Mount Mulu and Mount Murud.

Ecology

Mostly in montane mossy forest but extending to open ridges, 1500–3500 m. Flowering: throughout the year.

Conservation assessment

LC. This species is widespread on the mountains of western Borneo, some of which have protected status. This is a common species on both Gunung Kinabalu and Gunung Mulu.

For comments on the differences between this species and Vaccinium simulans, see under the description of the latter.

Fig. 40.

Vaccinium phillippsiae Argent. Lamb & Phillipps 2205; Crocker Range, Sabah. (Scan: Robyn Drinkwater.)

Terrestrial shrub. Twigs minutely patently pubescent, leaves laxly arranged, lateral buds c.2 × 1.5 mm, very narrowly conical, inconspicuous, as long as or longer than subtending petioles, scales subulate, hairy. Leaves: petiole 1–2 × c.0.75 mm, minutely hairy, without a groove; blades 25–40 × 13–20 mm, elliptic to weakly obovate; base tapering, margin entire, narrowly revolute, fringed with minute hairs when young, with 1 or 2 marginal glands on each side 2–3 mm from petiole; lamina with brown glandular hairs beneath when young, minutely greyish pubescent above, apex shortly acuminate, tip narrowly acute, midvein smooth to weakly raised above, minutely but distinctly raised below, lateral veins 1 or 2, each side high-arching from near base and with several smaller veins arising distally, reticulation conspicuous on both sides. Inflorescence of solitary racemes in upper axils, flowers 4–7, secund; rachis 12–25 mm, laxly hairy with simple patent hairs; bracts and bracteoles, not seen. Flowers: pedicels 6–8 × 0.4 mm, minutely laxly patently hairy with simple hairs; calyx covered all over with long (to 0.2 mm), simple patent hairs, tube cup-shaped, c.12 × 20 mm, lobes triangular, c.1.2 × 1.2 mm, with an apical gland and several sessile marginal glands; corolla 8–10 × c.4 mm, narrowly conical, pink, glabrous outside, but spotted with grey wax outside when dry, laxly hairy proximally inside, lobes c.1 mm reflexed; stamens: filaments 2.5–4 mm, broad at base, narrowing upwards to become very slender, densely hairy to apex; anther cells c.0.6 mm, oblongoid, with 2 distinct awns, muriculate with short points, tubules c.1 mm, with small glandular hairs, pores circular, opening slightly obliquely; disc glabrous at base, densely long hairy in upper part; style c.9 mm, densely hairy in proximal 3/4, with distally pointing hairs, slightly expanded distally. Fruit not recorded.

Distribution

Indonesia: East Kalimantan, Mount Kemul. Malaysia: Sabah, Mount Kinabalu, Crocker Range, northern Sarawak.

Ecology

Montane forest, 1300–1770 m. Flowering: August to September.

Etymology

Named after the late Mrs Susan Mary Phillipps, keen naturalist and long-standing resident of Sabah, for her hospitality and help over many years.

Vaccinium phillippsiae is very distinctive with its small (to c.10 mm) leaves with acute apex, rigidly patent hairs on stems, inflorescence with muriculate anther cells and anther tubules as long as or longer than the cells. This species also occurs at higher elevation than Vaccinium bancanum. The description has been augmented from the Lamb & Phillipps specimen (see Fig. 40) which is excellent flowering material. The name ‘kemulense’ was pre-dated at specific rank and so, if used, would create a later homonym.

Fig. 41.

Vaccinium phillyreoides. Cultivated RBGE accession number 19801159; Crocker Range, Sabah. (Photograph: G. Argent.)

Mostly epiphytic, occasionally terrestrial shrub to 2 m. Twigs spreading, minutely hairy, subdensely to laxly leaved; lateral buds inconspicuous but present even on very young stems, up to 2 mm long, as long as or slightly longer than petioles, narrowly conical, scales with subulate points. Leaves: petiole c. 2 × 0.7 mm, puberulous, clearly grooved above, usually red when fresh; blade 22–30 × 5–10 mm, base broadly tapering to rounded, margin entire or weakly revolute, fringed with minute hairs when very young with one small marginal gland each side 2–3 mm above petiole, lamina minutely and sparsely glandular hairy beneath, otherwise glabrous, midvein strongly impressed above and raised beneath throughout its length, lateral veins and reticulation mostly obscure when fresh, 1–3 lateral veins per side visible when dry, proximal veins high-arching, distal ones short and straight, narrowly ovate to elliptic, apex long acuminate, extreme tip acute to rounded. Inflorescence of racemes from upper axils, sometimes terminal, flowers 4–12, secund; rachis 20–40 mm, red, shortly white-hairy; bracts up to 4 × 1.5 mm, elliptic, acuminate, fringed with small glands and a few simple hairs withering and falling well before anthesis; bracteoles to 1.5 × 0.3 mm, narrowly triangular, inserted just above slightly swollen base of pedicel, minutely hairy. Flowers: pedicels 5–7 mm, red, white-hairy; calyx tube, 1–1.5 mm, globose, hairy, limb spreading, lobes 1–1.5 mm, hairy along a broad middle line outside, with a marginal rows of small sessile glands and with a small or only slightly larger terminal gland; corolla 8–9 × c.2.5 mm, tubular-urceolate, slightly contracted distally, red to pink, scented of almonds, glabrous both outside and inside or with some hairs proximally inside; lobes c.1 mm, pale pink to almost white, reflexing; stamens: filaments slightly dimorphic, alternately 6 and 7 mm, filiform, with lax, long white hairs; anther cells 1 mm, oblongoid, without spurs; tubules 0.2–0.3 mm, as wide as cells, opening obliquely; disc densely white-hairy; style c.8 mm, columnar, densely hairy proximally, glabrous distally. Fruit c.5 mm in diameter, shortly pubescent, with incurved calyx lobes, ripening black.

Distribution

Brunei. Indonesia. Malaysia: Sabah and Sarawak. Widespread on mountains in Borneo.

Ecology

In montane rain forest, 700–1830 m. Flowering: October to March, probably occasionally throughout the year. In cultivation in Scotland, flowering in late spring with occasional flowers at other times of the year.

Conservation assessment

LC. Common on many mountains, some with protected status.

A pretty and distinctive species with small leaves which have a deeply impressed midvein, obscure lateral veins when fresh, and long-acuminate apices.

Fig. 42.

Vaccinium retivenium. Mount Kinabalu, Sabah. (Photograph: G. Argent.)

Shrub or small tree to 8 m. Twigs glabrous with lax leaf arrangement; lateral buds up to 1.3 × 1.5 mm, often obscure, hemispherical, glabrous except for a minute fringe of hairs on scale margins. Leaves: petiole 10–20 × 1.5–2 mm, flattened and grooved above; blade 80–140 × 30–70 mm, ovate-elliptic; base broadly attenuate to rounded, extreme base shortly and abruptly contracted, margin flat, entire, with 2–4 marginal glands distributed along proximal half of blade, apex for 15–30 mm caudate-acuminate, acutely pointed; midvein prominent on both sides, with 3 or 4 prominent ascending veins also raised on both sides. Inflorescence of racemes from upper leaf axils, laxly 6- to 10-flowered, the flowers secund; rachis 50–100 mm, without bracts at flowering, glabrous, bracts and bracteoles not seen or recorded. Flowers: pedicels 13–20 × c.1 mm, often curved and swollen distally, glabrous (longer and thicker in fruit); calyx tube 4–5 × 4 mm, narrowly cup-shaped to cylindrical with rounded base, lobes c.1 mm, broadly triangular, obtuse with thick apical gland; corolla c.6 × 5 mm, broadly urceolate to subcampanulate, white, thick and fleshy, glabrous on both sides; lobes c.2 mm, suberect; stamens: filaments c.1.8 mm, linear, densely hairy; anther cells 1.5 mm, oblongoid, shortly 2-spurred dorsally or without spurs; tubules 0.5 mm, nearly as wide as cells, a little incurved, pores oblique; disc low, glabrous; style c.5 mm, glabrous. Fruit subovoid-globose, apex truncate, calyx lobes inflexed, ripening black.

Distribution

Malaysia: Sabah, Mount Kinabalu, Mount Alab, Mount Trus Madi and Mount Lotung; Sarawak, Mount Kalulong and Mount Pagon.

Ecology

In montane mossy forest, 1000–2500 m. Flowering and fruiting: throughout the year.

Conservation assessment

LC. Although not known to be widely distributed, it is locally abundant and growing in protected areas.

This is a distinctive species with its long petioles and pedicels displaying a very lax habit. The bracts and bracteoles have not been described, as they are totally lacking from the herbarium specimens examined and apparently wither and fall extremely early in development of the inflorescence. A record of this species from East Kalimantan (Beaman et al., Reference Beaman, Anderson and Beaman2001) remains very doubtful, since the collection was made far from its otherwise restricted northwesterly distribution.

17a. Vaccinium sarawakense subsp. sarawakense. Figs 43, 44.

Fig. 43.

Vaccinium sarawakense subsp. sarawakense. RBGE accession number 19851909. A, Flower (scale bar, 5 mm); B, bract (scale bar, 5 mm); C, stamens and style with disc (scale bar, 5 mm); D, young inflorescence (scale bar, 5 mm); E, habit (scale bar, 5 cm). (Illustration: Claire Banks.)

Fig. 44.

Vaccinium sarawakense subsp. sarawakense. Cultivated RBGE accession number 19851908; Nabawan, Sabah. (Photograph: G. Argent.)

Shrub, climbing to c.7 m. Twigs rounded, densely covered with appressed glandular hairs which are at first transparent, later turning brown, also with minute patent hairs, becoming glabrescent, laxly leaved; lateral buds mostly obscure, occasionally prominent, projecting from defoliate axils, hemispherical, up to 1 mm in diameter, scales broader than long with a few glandular hairs abaxially and with a minute glandular fringe on margins. Leaves: petiole 4–6 × c.2 mm, shallowly grooved, glandular hairy when young; blade 70–110 × 25–50 mm, elliptic to broadly elliptic, occasionally ovate, base broadly to narrowly attenuate, margin narrowly cartilaginous, entire, flat, marginal glands a single pair at c.1 mm from petiole, often slightly protruding, when young, lamina with appressed glandular hairs both above and below, midvein impressed above and prominent below, lateral veins 2 or 3 per side spreading at an acute angle, one pair from just above base ascending into upper part of leaf but disappearing before margin, upper pair shorter but similar, reticulation obscure, apex shortly attenuate, acute, extreme tip sometimes rounded. Inflorescence of racemes from upper axils, occasionally from defoliate axils, flowers 6–12, secund; rachis 25–50 mm, rounded, densely appressed glandular-hairy; bracts 6 × 2.5 mm, glabrous with indistinct midvein and cucullate apex; bracteoles minute, c.0.4 mm, broadly ovate with terminal gland. Flowers: pedicels 2.5–4 mm, less densely brown glandular-hairy than rachis; calyx tube c.1 × 2.4 mm, hemispherical, with just a few glandular hairs, limb erect, deeply lobed, lobes 1.5 × 1.6 mm, triangular, glabrous with no marginal glands or hairs but a faint translucent cartilaginous border, with a conspicuous but small, terminal gland; corolla tubular-urceolate 7–8 × 4–5 mm, glabrous outside, sparsely hairy inside, white or pink, scented; lobes c.1 × 1.5 mm, ovate, recurved; stamens: filaments c.2.3 mm, subulate, glabrous distally, long-hairy and broadening at base where adherent to disc; anther cells 0.8 × 0.6 mm, oblongoid, with short spurs, tubules c.1.2 mm, slightly narrower than cells, truncate, pore circular, margin reflexing, without gland-tipped hairs; disc glabrous; style c.6.5 mm, glabrous. Fruit not seen.

Distribution

Malaysia: Sabah, Nabawan; Sarawak, Kuching Division, near Kuching, Miri Division, Kelabit Highlands.

Ecology

Epiphytic in kerangas (heath forest) and along rivers, 300–600 m. This subspecies is a weak climber with arching stems rooting freely wherever they strike a suitable substrate. Flowering: March and May in the wild, March, December and January in cultivation at the Royal Botanic Garden Edinburgh.

Conservation assessment

It is suspected that the habitat of this species near Kuching has already been destroyed. In the Nabawan forest, it is not a common plant and this area may also be under threat of development, although the poor, acid nature of the soil may protect it.

This subspecies is the only member of Vaccinium sect. Bracteata which might be mistaken for a species of Rigiolepis due to its climbing habit. It has not been reported with a basal woody tuber, however, nor does it have large lenticels and the flowers are completely consistent with Vaccinium sect. Bracteata. See further discussion of this species below.

Fig. 45.

Vaccinium sarawakense subsp. montanum Argent. Type, Davies et al. 99145. Crocker Range, Sabah. (Scan: Robyn Drinkwater.)

Free-standing shrub to 2 m or small tree. Twigs robust, 3–5 mm in diameter, rounded or angular, indumentum of short, curved, brown glandular hairs which are quickly eroded, stems then glabrescent, laxly leafy; lateral buds 2–3 mm, rounded on young stems, sometimes narrowly pointed, often standing out at right angles on older, leafless portions. Leaves: petiole c.5–6 × 2 mm, scabrous, at first with glandular hairs, grooved when dry; blade 70–120 × 40–50 mm, elliptic, base tapering and shortly decurrent, midvein impressed above, strongly raised beneath, lateral veins 3–5, arising in proximal half of leaf, ascending to upper part of leaf, slightly raised above, reticulation obscure or somewhat distinct above, margin, entire, flat or only narrowly revolute, apex acute. Inflorescence of racemes from leafy axils, 7- to 20-flowered; rachis 60–80 mm long, at first densely covered in short brown curved glands, without simple hairs; bracts and bracteoles not recorded. Flowers: pedicels 2–3 mm, glandular hairy; calyx tube c.1 × 1 mm, hemispherical, densely covered with brown glandular hairs, 5-lobed, lobes c.1 × 1 mm, triangular, glandular hairy abaxially, and ciliate along margins, with simple hairs, lacking both lateral and terminal glands; corolla c.10 × 3 mm, pale pink, musty smelling in the evening, ovoid-urceolate, inflated below, somewhat contracted distally, glabrous outside and inside; lobes up to 1 mm, becoming reflexed; stamens: filaments 3 mm, linear, long-hairy at base; anthers c.0.8 mm, cells broadly oblongoid, echinulate, with distinct spurs to 0.3 mm, tubules 10 mm, narrower than cells, opening by slightly oblique, round pores, without teeth; disc glabrous; style to 8.5 mm, filiform, glabrous. Fruit not recorded.

Distribution

Malaysia: Sabah and Sarawak.

Ecology

Submontane forests recorded both terrestrial and epiphytic. (900–)1200– 1500 m. Flowering: October to November.

Conservation assessment

LC. Although there are no data on the specimens indicating abundance or frequency, this subspecies has been collected from well-protected forests on Kinabalu and the Crocker Range.

The essential differences between Vaccinium laurifolium var. laurifolium and V. laurifolium var. sarawakense, according to Sleumer (Reference Sleumer1966–1967), are the presence of numerous short brown glandular hairs on the rachis (and sometimes on the pedicels) and the presence of glands on the apices of the calyx lobes. These glands are consistently present in the Nabawan plant but can also be found on the Javan specimens (they are not mentioned in Sleumer's description), although their absence is used in his key to the varieties (Sleumer, Reference Sleumer1966–1967). As far as can be told from the specimens examined, subspecies sarawakense has apical glands on the calyx lobes and subspecies montanum does not. The biggest difference noticed both in the wild and in cultivation is that the Nabawan plant is a slender climber, as opposed to both the Javan plants, and apparently subspecies montanum, which are rigidly robust erect shrubs or trees. There are minor differences in the leaves, subspecies sarawakense having leaves tapering to an attenuate base with a pair of glands on the margin just above the junction with the petiole. The Javan specimens have decurrent bases to the leaves with the basal glands on the decurrent portions of the leaf blades and so these appear almost as if on the petioles, and this structure is very similar to that which occurs in subspecies montanum. The petioles themselves are about twice as long as broad in the Nabawan specimens but often more than 3 times as long as broad in the Javan specimens which is again similar to subspecies montanum. The axillary resting buds (not described by Sleumer, Reference Sleumer1966–1967) are quite different in the geographical forms, the Javan plants (Javan Vaccinium laurifolium) having shortly conical buds to c.2 mm long with acute apices and the Nabawan plant having rounded, hemispherical buds less than 1 mm long, while subspecies montanum has mostly rounded buds on young stems occurring on the older leafless portions, but the buds can be narrowly pointed, often characteristically standing out at right angles in sterile specimens. The anthers of Vaccinium sarawakense and V. laurifolium also differ, the tubules being shorter than the cells in V. laurifolium (cells 1–1.5 mm, tubules 0.5–0.8 mm; Sleumer, Reference Sleumer1966–1967) versus cells 0.9 mm, tubules 1.2 mm (RBGE accession number 19851909), i.e. distinctly longer than the cells; they also only flare at the extreme apex and have not been observed with the tooth-like extensions so common in V. laurifolium on other islands. There would also appear to be a distinct difference in the ecology of taxa in this complex, Vaccinium laurifolium being a montane plant “flowering abundantly between 2000 and 3000 m” (Sleumer, Reference Sleumer1966–1967) although recorded sterile as low as 800 m. The Nabawan plant (subspecies sarawakense) was fully fertile in the podzol forest at 600 m, and this is considered to be a lowland subspecies, whereas subspecies montanum has been collected flowering at around 1500 m and would appear to be a montane plant like Javan Vaccinium laurifolium.

It is felt that both Bornean forms are best kept separate from Vaccinium laurifolium and maintained as subspecies at least until much more is known about this complex. The Bornean subspecies share the brown glandular indumentum on the inflorescences and very similar anther structure. The status of other varieties of Vaccinium laurifolium which are outside the scope of this study are open to question, and I have presently excluded Vaccinium laurifolium (Blume) Miq. var. glanduligerum Sleumer as a Sumatran form of this complex, the type (Steenis 9962; BO, K, L), having been collected in Aceh Province at the confluence of the rivers Kapi and Aunan. There are still problems in interpreting what may be termed the Vaccinium laurifolium complex, and further observations are certainly needed. Merrill's name Vaccinium sarawakense has been resurrected after comparing herbarium specimens and especially the living material growing at the Royal Botanic Garden Edinburgh, where we have V. laurifolium var. laurifolium cultivated from Javan material (RBGE accession numbers 19680822 and 19680858) and a collection that keyed in Sleumer (Reference Sleumer1966–1967) to V. laurifolium var. glanduligerum from Nabawan in Sabah (RBGE accession number 19851909).

Distribution

Indonesia, West Kalimantan. Malaysia: Sabah, Mount Kinabalu.

Ecology

In primary ridge forest, terrestrial or epiphytic, sometimes recorded as a strangling climber, 1220–2000 m elevation. Common. Flowering: March to June.

Conservation assessment

LC. Well protected in the montane forests of Gunung Kinabalu National Park.

A distinctive species with the abaxial surface of the leaves smooth and only the midvein raised, in contrast to Vaccinium pachydermum, which has both midvein and lateral veins distinctly raised abaxially. See further comments below under Vaccinium simulans var. hirtirachis. In Flora Malesiana (Sleumer, Reference Sleumer1966–1967), this species is described as a Kinabalu endemic. It is now known to be much more widely distributed. A sterile specimen Keßler et al. PK 2851 (E) from East Kalimantan, plateau of Mount Meratus, agrees well but is very isolated from other localities and needs better material for reliable evaluation. This Keßler specimen could also be another record of Vaccinium ceraceum, although the leaves are larger. There are also specimens collected on the peak of Balikpapan (East Kalimantan) (Kostermans 7335 and 7605) which agree well vegetatively, but there are no flowers and the fruits are galled. Descriptions on herbarium sheets reporting this species as a ‘strangler’ are highly doubtful. It is unlikely to be truly strangling when growing epiphytically.

Fig. 46.

Vaccinium simulans var. hirtirachis Argent. Type; Marai Parai, Mount Kinabalu, Sabah. (Scan: Robyn Drinkwater.)

Bracts green, elliptic, concave, up to 10 × 4 mm, with appressed brown glandular hairs abaxially and a fringe of short white hairs near the apex, otherwise glabrous. Rachis distinctly hairy; pedicels to 15 mm, laxly to densely hairy.

Distribution

Malaysia: Sabah, Gunung Kinabalu only; Sarawak.

Ecology

Growing together with the type variety. Flowering: February and March.

On morphological grounds, Vaccinium simulans would appear to be closely related to Vaccinium loranthifolium Ridl. from Peninsular Malaysia. It is somewhat doubtful whether this is really a strangling climber rather than just an epiphyte as recorded on some specimens. Vaccinium simulans Sleumer and Vaccinium pachydermum Stapf can be difficult to distinguish. In Vaccinium simulans, the pedicels are rarely > 3 mm long (except for var. leptopodum and var. hirtirachis known from Gunung Kinabalu and Sarawak), the lateral veins viewed from the underside of the leaf are hardly raised and often obscure, and the young stems are glabrous. In Vaccinium pachydermum, the pedicels are > 5 mm long, the lateral veins viewed from the underside of the leaf are distinct and clearly raised, and the young stems are pubescent, this pubescence often turning black with contamination of other organisms.

Flowers: pedicels more slender than in the type, 3–6 mm at flowering; calyx smaller than in the type variety, the limb lower more shortly 5-lobed or merely undulate; corolla 5 mm long.

Distribution

Malaysia: Sabah, Gunung Kinabalu only, growing with the type variety from 1065–1585 m.

Ecology

As for the type variety. Flowering: June.

Conservation assessment

LC. Protected as for the type variety.

Shrub or small tree up to 5 m. Twigs slender, glabrous or minutely hairy, leaves laxly arranged; lateral buds minute, up to c.1 mm, conical, less than half the length of subtending petioles. Leaves: petiole 2–3 × c.1 mm, rugulose, glabrous; blade 35–60 × 15–40 mm, obovate to elliptic, base broadly tapering, margin slightly revolute especially towards base, with 1 or 2 minute impressed marginal glands on each side just above petiole, lamina glabrous except for lax glandular hairs underneath, apex broadly obtuse to rounded, midvein weakly impressed above in proximal half and minutely raised below throughout its length, with 2 pairs of high-arching veins from near base and often a further pair emanating from above middle of leaf, all veins slightly raised above, more prominently so beneath, reticulation finely distinct, minutely raised on both sides when dry. Inflorescence of lax racemes from distal axils, flowers 4–8; rachis 20–40 mm, slender, angular, laxly patently hairy and with minute globular brown glands; bracts and bracteoles not seen or described, deciduous before flowering. Flowers: pedicels 3–4 mm, with patent hairs and glands as on rachis; calyx densely hairy and with a few subspherical brown glands, tube 1.5–2 mm, hemispherical, lobes ovate-triangular, subacute to obtuse, terminated by a thick apical gland and with several lateral glands on each side in addition to ciliate hairs; corolla narrowly tubular, 6–8 × 2.5–3 mm, white, fragrant, glabrous outside, densely hairy proximally inside; lobes c.1 mm, obtuse, becoming reflexed; stamens: filaments c.1 mm, linear, densely hairy throughout; anther cells c.0.4 mm, broadly oblongoid, echinulate, with dorsal spurs to 0.2 mm; tubules c.0.2 mm, cylindrical, slightly narrower than cells, and held at an angle to them, with several stalked or subsessile gland-tipped hairs, pores round, transverse; disc densely appressed hairy; style c.8 mm, hairy almost to apex. Fruit globose, c.5–6 mm in diameter.

Distribution

Brunei. Malaysia: Sarawak.

Ecology

Occurring at low elevation from 180 m to possibly 1000 m. In sandy, acid habitats, mostly terrestrial. Flowering: January and June.

Sleumer (Reference Sleumer1963) lists several specimens which have not been checked. I have modified the description with characters from Brunig S.10410, which has good flowers but agrees in most respects with the species as described by Sleumer (Reference Sleumer1966–1967). Apart from the obviously erroneous measurements of the anthers, the main difference from the type description is the dense hairs inside the corolla. Superficially Brunig S.10410 looks similar to Vaccinium clementis but the lowland habitat is quite different to that montane species and the axillary buds are much shorter than the petioles subtending them, as in V. bancanum. From Vaccinium bancanum, this species differs in the coarsely hairy calyx tube, style hairy almost to the stigma and the shorter anther tubules.