OLD WORLD GESNERIACEAE X: BORNEAN SPECIES OF CYRTANDRA ALLIED TO C. PHOENICOLASIA

Cyrtandra phoenicolasia , C. hottae and five new species constitute a group distinguished by a suite of characters including setose vegetative parts, deeply divided calyx, small white corolla marked on the lower lip with yellow to brown, unilateral disc, ovary constricted at base, and fruit small. All species are fully described and an identification key provided.


INTRODUCTION
The group of species closely allied to Cyrtandra phoenicolasia Lauterb. and C. hottae B.L.Burtt share the following characteristics: stem simple above the base, leafy towards the apex (lower leaves shed), often floriferous well below the leaves, conspicuous brown bristles (setae) on all vegetative parts, inflorescence an axillary tightly congested, dichasial cyme, peduncle not exceeding c.35mm, mostly much shorter (3-10mm), calyx deeply divided (roughly halfway in C. phoenicolasia, otherwise nearly to the base), corolla small (c.12-20mm), white marked with yellow to brown on the lower lip, ovary glabrous, constricted at base on ventral side to accommodate the fleshy unilateral disc, style puberulous or minutely gland-dotted, fruit small (c.5-15mm long), pericarp verrucose.
In the shortly pedunculate, very congested inflorescence, deeply divided calyx, small white corolla, ovary constricted at the base to accommodate the unilateral disc, and in their small fruits, C. phoenicolasia and its allies resemble C. chrysea and its allies (Hilliard & Burtt, 2004). The latter group, although variously hairy, differs in its lack of brown bristles and also in the pink to purple (not yellow to brown) markings on the corolla.

Key to species
1a. Leaves with two conspicuous balloon-like pouches c.7-8mm in diam. at each node formed by the united bases of the petiolar wings --1. C. phoenicolasia 1b. Leaves without such pouches ----------------------2 2a. Hairs on stems closely appressed; largest major leaves c.37-45mm broad, minor leaf roughly one-tenth to one-third length of major leaf ----7. C. hottae 2b. Hairs on stems spreading; largest major leaves 50-150mm broad; if leaves anisophyllous, then minor leaf one-third to two-thirds length of major leaf -3  Herb, stem to c.600mm long, 4-5mm diam., simple, decumbent at base and rooting there, thickly clad in brown bristles c.3-5mm long. Leaves opposite, anisophyllous, minor leaf roughly a third as long to half as long as major leaf, largest major leaves 120-200x55-85mm, obovate, apex abruptly acute, base cuneate, decurrent, margins serrate to serrulate, each tooth a vein-ending, lateral veins 10-12 each side of midrib, tertiary veins coarsely reticulate, all veins strongly raised on lower surface, upper surface clad in scattered brown bristles 3-5mm long, c.2mm on margins, a small 'shield' (outgrowth of blade) c.6-7.5x5-7.5mm at base of midrib, apical half free of midrib, lower surface glabrous between the veins, all veins and veinlets densely appressed-pubescent, bristly as well, bristles to 3-4mm long; petiole c.10-15mm long, winged by decurrent blade, wings broadening at base and united with wings of opposing leaf to form a deep pouch c.7-8x7-8mm. Inflorescence a dichasial cyme, solitary in leaf axils, flowers c.10, tightly clustered; peduncle c.8-10mm, densely bristly. Bracts c.13-17x2-8mm, sublinear to elliptic, acuminate, venation clearly visible particularly on lower surface, brown bristles to c.4mm on margins, fewer on inner and outer surfaces, bracteoles similar but narrower. Pedicels 2-4mm long, bristly. Calyx 2-3mm long, campanulate, tube 1-2mm, lobes 5, 1-1.5x0.8-1mm, bristly to glabrous. Corolla white with 2 raised yellow keels on lower lip running down into throat, c.14-17mm long, tube 9.5-12mm, cylindric at base, then abruptly expanded and ventricose on lower side, bilabiate, lower lip 4.5-5x7-9mm, median lobe 2. A remarkable feature of C. phoenicolasia is the two deep pouches at each node formed by the union of the winged bases of the petioles of each pair of leaves. In addition, there is a shield-shaped structure on the upper surface of the leaf astride the midrib near its base. This appears to be an outgrowth of the blade, attached to the midrib only in its lower half. What purpose it serves is unknown; the pouches are possibly 'compost pits', though in one specimen seen (Anderson S28640) they were filled with small seeds, perhaps cached there by ants. Union of winged petioles to form 'pockets' has been recorded in several other, unrelated, species of Cyrtandra, but, so far as is known, never in the form recorded here.
Specimens from Kalimantan appear to have bracts broader than in those from Sarawak, but this needs confirmation on more plentiful material.
A broad concept of C. phoenicolasia has hitherto included a number of allied species newly described below as C. brevisetosa, C. dorytricha, C. latibracteata, C. megaphylla and C. phoenicoides. All lack the leaf pouches of C. phoenicolasia and differ further in several respects, including a more deeply divided calyx.
Plants of C. phoenicolasia favour outcrops, boulders and cliffs in forest along streams, from SE Sarawak eastwards across Kalimantan, between 300 and 850m above sea level.
The leaves lack a hypodermis and there are no sclereids in the mesophyll. Herb, stem to 1m tall, probably decumbent at base (adventitious roots present), c.7-10mm diam., thickly clad in brown bristles 7-12mm long, leafy only towards apex. Leaves opposite, anisophyllous, minor leaf roughly one-half to two-thirds as long as major leaf, largest major leaves 270-370x88-150mm, including petiolar part, obovate, apex rather abruptly acute, base cuneate, decurrent down very short petiole in wings c.8-10mm broad, wings of leaf pairs briefly united at base, margins serrulate, each tooth a vein-ending, lateral veins 13-20 each side of midrib, tertiary veins coarsely reticulate, often scalariform towards margins and apex, upper surface clad in scattered bristles 3-4mm long, lower surface glabrous between the veins, all veins and veinlets densely appressed-pubescent, spreading brown bristles 8-10mm long as well, longest on midrib. Inflorescence a many-flowered very congested, axillary, dichasial cyme; peduncles 15-34mm long, clad in brown bristles c.4mm long. Bracts c.18-30x3-5mm, lanceolate, acuminate, venation obscure, brown bristles to c.5mm long on both surfaces, bracteoles similar but narrower and slightly shorter. Pedicels c.2-5mm long, bristly. Calyx 5-lobed nearly to base, tube c.0.25-0.5mm long, lobes 2-2.5x0.8-1.2mm, narrowly triangular, bristles 1-2mm long outside. Corolla white with 2 yellow bars on palate darkening to brown in throat, c.19mm long, tube 13mm, cylindric at base, then abruptly expanded and ventricose on lower side, bilabiate, lower lip c.6x11mm, median lobe 3x4.5-5.5mm, upper lip 3x7-8mm, lobes 3x4mm, outside brown appressed hairs to 1mm long, inside minute hairs on lower lip, particularly on the yellow bars. Stamens inserted 8mm above base of tube, filaments 5.5mm long, twisted once, anthers 1.5x1.25mm, cohering face to face by a very small apiculus; lateral staminodes 1mm long, posticous staminode 0. Cyrtandra megaphylla (megaphylla = large leaves), like the rest of the species in this group, differs sharply from C. phoenicolasia by the lack of pouches at the nodes derived from the united petiolar wings of the leaf-pairs and in having the calyx divided nearly to the base (not roughly halfway). It differs from all other species included here in its much longer peduncles, 15-35mm versus 3-10mm.

Cyrtandra megaphylla
The distribution of C. megaphylla as currently known is confined to Sarawak between 1°30p and 2°42pN and 113°5pS to 47pE, while that of C. phoenicolasia is further south and east, between 1°35pN to 1°50pS, 114°30pE to 116°2pE, from E Sarawak east across Kalimantan. Cyrtandra megaphylla is a plant of the forest floor along rocky streambanks and on hill slopes, c.500-700m above sea level.
Burtt recorded 'young foliage copper-purple underneath, gradually fading'; Yii 'stems and petioles purplish'. Others commented on the brown or blackish hairs on mature stems and leaves, a feature of this group (they also occur in other groups). The leaves have a one-layered hypodermis and small astrosclereids in the spongy mesophyll. Herb, stem c.250-500mm long, 4-5mm diam., decumbent and rooting at base, sometimes with a minor branch near base, thickly clad in brown bristles to 5mm long. Leaves opposite, more or less isophyllous to anisophyllous and then minor leaf roughly half length of major leaf, largest major leaves 150-250x55-100mm long including petiolar part, elliptic to obovate-elliptic, apex acute, base cuneate, decurrent down short petiole in wings c.3-4mm broad, wings of leaf-pairs briefly united at base, margins serrate, lateral veins 11-14 each side of midrib, tertiary veins coarsely reticulate tending to scalariform towards margins, upper surface with scattered brown bristles 2.5-5.5mm long, lower surface glabrous between the veins, all veins and veinlets densely appressed-pubescent, brown bristles 4-5.5mm as well mainly on midrib and lateral veins. Inflorescence a very congested axillary, dichasial cyme, flowers 10 to many; peduncles 3-5mm long, bristly. Bracts 12-16x2-3mm, lanceolate, acuminate, venation invisible, brown bristles to 3mm long mainly on backs and margins, bracteoles similar but narrower. Pedicels 2-7mm long, bristly. Calyx 5-lobed nearly to base, tube c.0.3mm, lobes 2.25-2.5x0.8mm, narrowly triangular, few small bristles outside. Corolla white with 2 yellow bars on palate turning brownish in throat, c.15mm long, tube 10mm, cylindric at base then abruptly expanded and ventricose on lower side, bilabiate, lower lip 5x10mm, median lobe 2.5x4mm, upper lip 2x4mm, notched, outside brown bristles 1.5mm long, inside minutely glandular below sinus of upper lip. Stamens inserted c.6mm above base of tube, filaments 4mm, twisted once, anthers 1x1mm, cohering face to face by a very small apiculus; staminodes not seen. Disc 1x1mm, unilateral. Ovary 3x0.8mm, constricted at base to accommodate disc, glabrous. Style 4mm, very minutely glanddotted. Stigmatic lobes 0.6x0.4mm. Fruit 5-6x3-3.5mm, pericarp verrucose. Seeds c.0.25x0.2mm. Cyrtandra phoenicoides is in the general affinity of C. phoenicolasia; the epithet draws attention to this. Its closest ally appears to be C. megaphylla, from which it is distinguished by its smaller major leaves (c.150-250mm versus 270-370mm long), shorter bristles on stem and undersurfaces of leaves (up to 5mm versus 7-12mm) and shorter peduncles (c.3-5mm versus 15-35mm). Also the bracts are shorter and narrower, and the flowers are possibly smaller, but too few corollas have been seen to be certain.

Cyrtandra phoenicoides
The two species are well separated geographically, C. megaphylla apparently confined to Sarawak south of 3°N latitude, while C. phoenicoides is currently known from Brunei and just across the border in Mulu National Park, c.4-5°N latitude.

Cyrtandra latibracteata
Other specimens examined. SABAH. Mt. Kinabalu, eastern shoulder, 6°05pN, 116°36-40pE, Singh's Plateau, 3000ft, 13 vi 1961, Chew et al. 1024 Cyrtandra latibracteata resembles C. phoenicolasia in its conspicuously veined bracts, but lacks petiolar pouches at the nodes, which are a conspicuous feature of that species. It differs further in its larger leaves, bigger bracts, and in the calyx being lobed nearly to the base. Its closest ally is possibly C. dorytricha, from which it is distinguished by its much bigger, strongly veined, bracts, longer calyx lobes, and possibly longer fruits. That we have seen no corollas of this species is of little importance; it is features of calyx and gynoecium that are of diagnostic value in this group.
Cyrtandra latibracteata occurs on Mt. Kinabalu, between c.900 and 1000m above sea level, in forest. Then there appears to be a record from 3° further south in Sarawak (Hotta 15418, cited above). This specimen differs from the Kinabalu plant by the presence of long bristles on both the midrib and lateral veins on the lower leaf surface. There is an urgent need for collecting before these forests are completely destroyed by logging.

Cyrtandra dorytricha
The epithet dorytricha (meaning spear hair) draws attention to the coarse brown bristles that are a characteristic of all the species in this group. Astrosclereids occur in the spongy mesophyll of the leaves; a hypodermis may be present or absent.
Cyrtandra brevisetosa is known to us only from the type collection. The plants were growing in shade among sandstone rocks by a river in the Gunong Bunga Range southwest of Kuching. The species has a distinct facies: long slender stems glabrous except at the apex and leafy there, with few inflorescences scattered down the stem well below the leaves.
The bristles on the stems are distinctly shorter than those of any other species in the group, which character suggested the specific epithet. Herb, stem up to c.500mm tall, 4mm diam., woody, simple, erect, leafy towards apex, often floriferous well down leafless part, thickly clad in strongly appressed brown bristles. Leaves opposite, strongly anisophyllous, minor leaf c.13-40x 4-12mm, roughly one-tenth to one-third size of major leaf, largest major leaves 130-190x37-45mm including petiolar part, narrowly obovate, abruptly acuminate, base cuneate, decurrent down very short petiolar part, margins crenate-serrate, both surfaces clad in red-brown bristles to c.3-5mm long. Inflorescence a 12-to manyflowered, very congested, axillary, dichasial cyme; peduncles c.5-8mm long, bristly. Bracts 12-18x6-9mm, ovate, bristles to 3mm on both surfaces mainly towards margins and on prominent midrib on lower surface; bracteoles similar, scarcely smaller. Pedicels 2-3mm long, setose. Calyx 5-lobed almost to base, lobes c.1.8-2x0.7mm, narrowly triangular, very few bristles on back of each lobe. Corolla white with a small patch of dark blotches (seen in dried material: no notes on colour by collectors) below the two sinuses of the lower lip, c.13mm long, tube 11mm, cylindric below then abruptly expanded and ventricose on lower side, bilabiate, lower lip c.2x6mm, median lobe 2x2.2mm, upper lobes 1.5x2.6mm, outside a few bristly hairs, inside glabrous. Stamens inserted 7mm above base of tube, filaments c.5mm long, slightly twisted, anthers 1x0.8mm, face to face and united at extreme tips, staminodes not seen. Disc c.1x1mm, unilateral. Ovary 2.5x1mm, constricted at base to accommodate style, glabrous. Style c.2mm, few minute glandular hairs towards base. Stigmatic lobes c.0.25mm long. Fruit 5x2mm, pericarp verrucose. Seeds not fully ripe. Cyrtandra hottae is rendered distinctive by the strongly appressed bristles on the stem in contrast to the spreading bristles in the other species of this group. Also, the leaves are often remarkably anisophyllous, and the bracteoles are scarcely smaller than the bracts, giving the inflorescence a very leafy look. Hansen described the stems and undersurface of the leaves as dark purple; the colour is often detectable in dried leaves. The leaves lack sclereids and a hypodermis.

Cyrtandra hottae
Dr Mitsuru Hotta, Research Associate in the Herbarium, Department of Botany, Kyoto University, collected Bornean plants in the 1960s and donated many duplicates to E.
No further collections have been made since the species was first described in 1990. The plant may have a narrow distribution within Sarawak, between roughly 2°40p and 3°N, 112°50p to 113°E.

A C K N O W L E D G E M E N T S
We are indebted, as ever, to the curators of the following herbaria for sending specimens on loan: The Natural History Museum, London (BM), Botanical Museum, University of Copenhagen (C), Harvard University Herbaria (GH), Royal Botanic Gardens, Kew (K), Botany Department, Kyoto University (KYO), Nationaal Herbarium Nederland, Leiden (L), Botanic Gardens Singapore (SING) and Botany Department, Museum of Natural History, Wroclaw University, Wroclaw (WRSL).
Prof. M.H. Bokhari (Multan, Pakistan) provided the information on leaf anatomy. We sincerely thank Prof. Douglas Henderson for putting our manuscript on disc.