TAXONOMIC REVISION OF BEILSCHMIEDIA ( LAURACEAE ) IN BORNEO

A revision of Beilschmiedia Nees (Lauraceae) in Borneo is given. Descriptions, distribution maps, illustrations of leaves, terminal buds, and flowers, and a key to the species are provided. Twenty-six species were recognized, including one newly described species. The new species is distinguished from the other Beilschmiedia species of Borneo in having pubescent anther apices. Beilschmiedia reticulata was excluded from the revision as an imperfectly known species because of a lack of flowering specimens. Beilschmiedia eusideroxylocarpa was also excluded because it had only six stamens representing the second and third whorls, which is characteristic for the genus Endiandra.


INTRODUCTION
Beilschmiedia Nees is one of the larger genera of the Lauraceae with approximately 250 species (Nishida 2001), and is best represented in tropical Asia and Africa (Van der Werff 2003). The genus comprises trees and rarely shrubs and is usually distinguished from other genera of the Lauraceae by the following characteristics: paniculate or racemose inflorescences that are not strictly cymose at the terminal division, bisexual and trimerous flowers with six equal to subequal tepals, six to nine fertile stamens representing the outer two or three whorls, two-celled anthers, and fruits lacking cupules. It is placed in the tribe Cryptocaryeae Nees together with other core genera such as Cryptocarya R.Br., Endiandra R.Br., and Potameia Thouars based on wood and bark anatomy and inflorescence structure (Van der Werff & Richter 1996). This classification is also supported by a molecular study (Chanderbali et al. 2001).
The genus Beilschmiedia was originally established by Nees to accommodate two Indian species: B. roxburghiana Nees and B. fagifolia Nees. Since then approximately 350 names have been published in the genus (Nishida 2001). Revisional studies of the genus have been made in several regions, such as China, Madagascar, Australia, and the Neotropics, but never in the Malesian regions except for the treatment by Kochummen (1989) in the Tree Flora of Peninsular Malaya. Borneo is famous for its high plant diversity and endemism, and I have focused on the species of this island as a precursor to a treatment of the genus in Malesia.

MATERIAL AND METHODS
Herbarium collections from A, BO, K, KEP, L, SAN, and SAR were examined. All dimensions given are for dried material except for the floral characters. Flowers were soaked in boiling water before observation and measurement. Trees or rarely shrubs. Leaves alternate, subopposite, or opposite, rarely clustered, pinnately veined. Inflorescences axillary, racemose or paniculate with terminal branches of panicles not strictly cymose. Flowers bisexual. Tepals 6, equal or subequal. Stamens 9 or 6, representing outer two or three whorls; anthers 2-celled, introrse or latrorse in first and second whorls and latrorse or extrorse in third whorl. Staminodia 3, 6 (when stamens 9 or 6, respectively), or absent. Pistil stylocarpellous; ovary superior, with single ovule. Fruits ellipsoid, pyriform, spherical, or spindle-shaped, lacking cupule.

BEILSCHMIEDIA
Distribution -About 250 species, pantropical, with the centre of diversity in Southeast Asia or Africa.

Habit
All the Beilschmiedia species of Borneo are trees, ranging in height from 8-40 m.

Indument
The indument types and pubescence observed among the Beilschmiedia species of Borneo show a great deal of variation, which can be used to delimit species.
For example, the vegetative parts of the plants vary from completely glabrous to densely pubescent, whereas pubescence is consistently observed on at least some portion of the inflorescences, such as on the rachis, tepals, or filaments. Those species with opposite leaves lack pubescence on most of their vegetative parts, except for B. dictyoneura Kosterm. and B. montanoides Kosterm. Subopposite-or alternateleaved species, however, usually have some hairs on at least their terminal buds. The only exception to this is B. glauca S.K.Lee & L.F.Lau, which has alternate leaves but is glabrous on all of its vegetative parts. Pubescence on the lower leaf surface is usually a stable character, but in some species, such as B. tawaensis Merr., the density of hairs can vary.
The hairs of the Beilschmiedia species of Borneo can be divided into three types based on their orientation (appressed, ascending, or erect), into two types based on their length (long or short), and into three types based on their straightness (straight, wavy, or curly). The minutely tomentulose species sometimes appear dusty to the unaided eye. The indument can vary on different parts of the same plant, while different indument types occur on the same part of a single plant in some species. However, the indument type or combination of types present on certain parts of a plant, particularly on the terminal buds, is usually consistent within a species, and can be used as a means of identification.
first and second whorls, not the second and third whorls as seen in Endiandra. In smallflowered species, the filaments in the first and second whorls are usually shorter than or as long as the anthers. Some, but not all, of the large-flowered species, however, have longer filaments even in the first and second whorls (e.g., B. gemmiflora, B. glabra, B. lucidula (Miq.) Kosterm., B. madang (Blume) Blume, B. oblonga Kosterm., and B. wieringae). The filaments in the third whorls are longer than or as long as the anthers in many of the species. The anthers are glabrous at the apex except for in a new species, B. telupidensis. The relative filament length in the first and second whorls and the pubescence on the anther apices can be used to delineate species.

Fruits
The fruits are often ellipsoid and around 3 cm long, but are sometimes spherical or spindle-shaped, and conspicuously large. Spherical fruits occur in B. crassa, B. kinabalu ensis, B. lucidula, B. micrantha, B. phoebeopsis, and B. tawaensis, whereas club-shaped (or sometimes ellipsoid) fruits occur in B. glauciphylla Kosterm., and spindle-shaped fruit is found in B. gynotrochioides Kosterm. The fruits of B. kinabaluensis, B. glauci phylla, and B. gynotrochioides are large (longer than 5 cm). The fruit pedicels are not usually thickened, but the thickened pedicels in such species as B. glabra and B. gyno trochioides obscure the border between the pedicel and fruit.
Sometimes specimens with relatively long oblong fruits of Endiandra are misidentified to Beilschmiedia in Borneo. Long oblong fruits with obtuse base and apex are rather rare for Beilschmiedia, but more common for Endiandra in Borneo and the Malay Peninsula. However, there are some exceptions in this tendency because both genera have a wide variation in fruit shape. As mentioned by Van der Werff (2001), fruiting specimens can often not be assigned to either genus with certainty because of their similarity in fruit and vegetative characters.

TAXONOMY OF BEILSCHMIEDIA IN BORNEO
I previously reported 28 species and one variety name on the Beilschmiedia specimens collected in Borneo (Nishida 2001). After a careful re-examination of the specimens, however, five species, i.e., B. curtisii Gamble, B. lumutensis Gamble, B. malaccensis (Meisn.). Hook.f., B. perakensis Gamble, and B. wallichiana (G.Don) Kosterm., were excluded from the Bornean flora; one species, i.e., B. eusideroxylocarpa Kosterm., was excluded from the genus; one species, i.e., B. reticulata Kosterm., was left as an imperfectly known species because of the lack of flowering specimens; and one species and one variety, i.e., B. oligocarpa Kosterm. and B. micrantha var. latifolia Merr., were transferred as synonyms of other taxa. Thus, three species have been described after the report and two species have been retained as independent species from the synonym list. As a result, 26 species are recognized in Borneo, including a new species.
The Beilschmiedia species of Borneo show a wide variation, which makes it difficult to adequately divide all the species into groups based solely on the morphology. However, at least for some of the species, several groups can be recognized, mainly by phyllotaxes, indument types, inflorescence or flower structure: These groupings are partly supported by cuticular characters (Nishida unpublished).
The relationships among the groups and with the other species are unknown. These groupings should remain informal until a comprehensive classification is available for the genus.

KEY TO THE SPECIES
The key to the species is mainly based on flowering material, because flowers are necessary to identify the genus. Fruit characters are added if they are useful, but are always accompanied by vegetative characters. A few species appear twice in the key because of variability in their characters. Small tree up to 15 m tall. Terminal buds narrowly ovoid, densely pubescent with long, erect, wavy, reddish brown hairs, and short, erect, curly, reddish brown hairs. Twigs terete, slightly angular when young, c. 2 mm diam., dark brown, pubescent with same type of hairs as on the terminal buds. Petioles 0.5-1 cm long, flat to slightly canaliculate above, drying dark brown, pubescent. Leaves alternate, evenly distributed, blade chartaceous, matt, elliptic or obovate, 7-11 by 3-5 cm, base cuneate, apex acute to slightly acuminate, margin flat, upper surface pubescent along the midrib with long wavy hairs, lower surface sparsely pubescent with erect hairs, slightly glaucous, midrib almost immersed above, raised below, secondary veins 7-9(-10) pairs, almost immersed above, raised below, tertiary veins not percurrent, minor venation fine, almost immersed above, slightly raised below. Inflorescences axillary, paniculate to racemose, 2-5 cm long, pubescent with long erect hairs, each bearing 5-30 flowers; floral pedicels of lateral divisions 1.5-2.5 mm long, pedicels of central flowers c. 3 mm long. Flow ers greenish white, c. 2 mm long, c. 2 mm diameter. Tepals 6, ovate, c. 1.5 by 1 mm, densely pubescent on both sides. Stamens 9, c. 1 mm long, filaments shorter than anthers in the first and second whorls, longer than anthers in the third whorl, filaments pubescent, stamens in third whorl with a pair of glands; anthers glabrous, obtuse at apex. Staminodes 3, sagittate. Pistil glabrous; ovary ovoid, c. 0.5 mm long; style terete, c. 0.6 mm long. Infructescences c. 10 cm long; pedicels c. 1 mm diam., thickened to c. 2 mm diam. below fruits, not constricted at apex. Fruits ellipsoid, c. 2 by 1 cm (immature), drying dark brown, surface smooth.  Distribution -Endemic to Borneo, known from E Sabah (Malaysia) and E Kalimantan (Indonesia).
Note -This species is distinguished from the other Beilschmiedia species of Borneo by its thick flower receptacle and thick, strongly coriaceous, narrowly ovate leaves. Flowers of B. crassa have a variation in stamen number from six to nine because stamens in the third whorl are often sterile. Trees to 30 m tall. All parts glabrous except for terminal buds, young twigs, and inflorescences. Terminal buds ovoid, pubescent with brown, short, appressed hairs. Twigs terete, slightly angular when young, c. 1.5 mm diam., dark brown, usually glabrous or sparsely pubescent with short appressed hairs when young. Petioles 1.5-1.7 cm long, canaliculate above, drying dark brown. Leaves opposite, rarely subopposite, evenly distributed, blade firmly chartaceous, narrowly elliptic, 8-16 by (2.5-)3-4.5(-6.5) cm; base cuneate to obtuse, apex acute, margin flat; upper surface often slightly glossy, lower surface glaucous; midrib impressed or slightly raised above, raised below; secondary veins 6-10 pairs, sometimes arching to the next vein and ending in a loop, slightly raised above, raised below; tertiaries not percurrent; minor venation relatively coarse, very slightly raised above, almost immersed below. Inflorescences axillary, paniculate, 3-4 cm long, sparsely pubescent with short erect hairs, each bearing 10-20 flowers; floral pedicels c. 2 mm long. Flowers greenish white, c. 1.5 mm long, c. 2.5 mm diameter. Tepals 6, almost equal or inner three tepals slightly smaller, broadly ovate, 0.5-1 by 1-1.7 mm. Stamens 6, representing first and second whorls, c. 0.7 mm long, filaments shorter than anthers, almost glabrous or pubescent only around the base; anthers glabrous, acute at apex. Staminodes 3-6 (3 in third whorl, 0-3 in fourth whorl), c. 0.5 mm long, narrowly deltoid, a pair of glands rarely present in third whorl, lacking in fourth whorl. Pistil glabrous; ovary roundish ellipsoid, c. 0.6 mm long; style terete, c. 0.5 mm long. Infructescences up to 7 cm long; fruit pedicels c. 1.5 mm diam., slightly thicker to c. 2 mm diam. below fruits, not constricted at apex. Fruits ellipsoid, c. 1.5 by 1 cm, drying dark brown, slightly warty. Distribution -Sabah (Malaysia), E, C and W Kalimantan (Indonesia). The species has also been collected from the Malay Peninsula and Sumatra.
Note -This is one of the rare Beilschmiedia species that have only six stamens. This characteristic sometimes occurs also in B. crassa, B. glauca, and B. kinabaluensis, but B. dictyoneura is easily distinguished from these three species by the terminal buds pubescent with appressed hairs and the opposite, chartaceous, narrowly elliptic leaves. ( Tree to c. 35 m tall. Terminal buds ovoid, light brown, pubescent with short, curly hairs. Twigs terete, angular when young, c. 3 mm diam., light brown, pubescent with light brown, short, curly hairs. Petioles 1.5-2 cm long, canaliculate or flat above, drying brown. Leaves alternate, relatively crowded around the terminal buds, blade chartaceous, elliptic, (7-)9-15(-17) by (3.5-)4-8 cm, base obtuse to cuneate, apex acute, margin slightly revolute, upper surface slightly glossy or sometimes matt, lower surface matt, glabrous, not glaucous, midrib slightly impressed or almost immersed above, raised below, secondary veins 8-10 pairs, slightly raised above, raised below; tertiary veins not percurrent; minor venation relatively fine, almost immersed or slightly raised above, slightly raised below. Inflorescences axillary, enveloped by an involucre when young, racemose to paniculate with little branching, 1-3 cm long, each bearing a few to 20 flowers; floral pedicels c. 1 mm long. Flowers greenish, c. 3 mm long, c. 3 mm diameter. Tepals 6, equal, oblong, c. 2.5 by 1 mm, outside pubescent, inside almost glabrous. Stamens 9, first and second whorls c. 1.3 mm long, third whorl c. 1.5 mm long, all filaments longer than anthers, pubescent, stamens in third whorl with a pair of glands; anthers glabrous, obtuse at apex. Staminodes 3, sagittate. Pistil almost glabrous; ovary ovoid, c. 0.5 mm long; style terete, c. 1.2 mm long. Infructescences 5-8 cm long; fruit pedicels c. 3 mm diam., not constricted at apex. Fruits ellipsoid, c. 3 by 1.7 cm, drying blackish brown, surface smooth. Distribution -W Sabah (Malaysia), E and C Kalimantan (Indonesia). This species is also distributed in Sumatra, Java, Celebes, and New Guinea.

Beilschmiedia gemmiflora
Habitat & Ecology -Primary forest on sandy soils or brownish soils. Altitude 10-600 m. Flowering and fruiting in August.
Note -Beilschmiedia gemmiflora is distinguished from the other species of Borneo by its inflorescences enveloped by involucres. This character can be recognized even in fruiting specimens because there are traces of involucral segments at the base of infructescences. Tree to 35 m tall. Terminal buds ovoid, dark brown, as thick as the twigs, glabrous or rarely sparsely pubescent only on the edge of buds or inside buds. Twigs terete, angular when young, c. 4 mm diam., greenish brown, glabrous. Petioles 1.5-2 cm long, canaliculate above, drying dark brown, glabrous. Leaves opposite, evenly distributed, blade relatively coriaceous, ovate to elliptic, 12-18(-26) by 5.5-7(-8) cm; base cuneate, apex acute, margin usually flat, both surfaces matt, glabrous, lower leaf surface not glaucous, midrib almost immersed above, raised below; secondary veins 6-9 pairs, almost immersed above, slightly raised below; tertiary veins not percurrent; minor venation relatively coarse, immersed above, very slightly raised below. Inflorescences axillary, racemose or paniculate with little branching, 2.5-4 cm long, each bearing a few to 30 flowers; floral pedicels of lateral divisions 3-8 mm long, pedicels of central flowers 5-10 mm long. Flowers yellowish or pale green, c. 5 mm long, c. 5 mm diameter. Tepals 6, equal, oblong, c. 3.5 by 1.5 mm, only inside pubescent. Stamens 9, c. 2.5 mm long, all filaments longer than anthers, pubescent, stamens in third whorl with a pair of glands; anthers glabrous, truncate at apex. Staminodes 3, sagittate. Pistil glabrous; ovary ovoid, c. 0.7 mm long; style terete, c. 1.5 mm long. Infructescences c. 10 cm long; fruit pedicels c. 15 mm diam., not constricted at the apex. Fruits ellipsoid with a pointed apex, 3-4.5 by c. 2.5 cm, drying brown, surface smooth or slightly rough. Distribution -In Sabah, E Sarawak (Malaysia), and E Kalimantan (Indonesia). This species has also been collected from the Malay Peninsula.

Beilschmiedia glauca
Distribution -In Borneo, Sarawak and Sabah (Malaysia), Brunei, and E Kalimantan (Indonesia). The species is also distributed widely from Hainan to the Malay Peninsula and N Sumatra.
Habitat & Ecology -Primary, lowland forest to hill forest. Altitude up to 870 m. Flowering: April; fruiting: July, September.
Note -Many Bornean specimens of this species were identified as "B. endiandrae folia" by Kostermans, but this name is a nomen nudum. As the nude name describes, this species has a finely reticulated venation pattern that is often seen in Endiandra. This character, alternate phyllotaxis, and the lack of pubescence except for the inside of the flowers, distinguish B. glauca from the other Beilschmiedia species of Borneo. Stamens 9, c. 1 mm long, filaments shorter than anthers in first and second whorl, longer than anthers in third whorl, filaments almost glabrous, stamens in third whorl with a pair of glands; anthers glabrous, obtuse at apex. Staminodes 3, sagittate. Pistil glabrous, ovary ovoid, c. 1 mm long; style terete, c. 0.5 mm long. Infructescences 8-15 cm long; fruit pedicels c. 7 mm diam., thickened to c. 10 mm diam. below fruits, not constricted at apex. Fruits club-shaped or broadly ellipsoid, c. 6.5 by 4 cm, drying dark brown, surface smooth. Distribution -Endemic to Borneo, known only from Sarawak (Malaysia).

Beilschmiedia glauciphylla
Habitat & Ecology -Mixed dipterocarp forest or heath forest on sandstone or basalt. Altitude up to 1000 m. Flowering: May, June; fruiting: April.
Note -I previously referred to B. glauciphylla as having club-or spindle-shaped fruits in terms of the difference between this species and B. microcarpa (Nishida 2006). This recognition included specimens identified as B. gynotrochioides which had been reduced as a taxonomic synonym of B. glauciphylla by Kostermans (1970). After re-examination of the specimens, however, I concluded that B. glauciphylla and B. gynotrochioides should be retained as different species, and that the fruit shape of B. glauciphylla should not be described as spindle-shaped, but club-shaped or ellipsoid. Beilschmiedia glauciphylla and B. gynotrochioides can be discriminated by the appearance of the minor venation (raised vs immersed on the upper leaf surface), and fruit shape (club-shaped or ellipsoid with rounded top vs spindle-shaped with pointed top). Differences between B. glauciphylla and B. microcarpa are recognized in leaf texture (coriaceous vs chartaceous), flower size (c. 4 mm diam. vs c. 2 mm diam.), and fruit size (c. 6.5 by 4 cm vs c. 3.5 by 1.6 cm). Tree to 18 m tall. Terminal buds narrowly ovoid, c. 2 mm diam., blackish brown, glabrous or rarely sparsely pubescent only on the edge of buds or inside buds. Twigs terete, compressed when young, 1.5-4 mm diam., dark brown to black, glabrous. Petioles 1.2 -1.6 cm long, flat to rounded above, drying dark brown, glabrous. Leaves opposite, evenly distributed, blade coriaceous, matt, elliptic, 10-16 by 4-6.5 cm, base obtuse, apex acute, margin slightly revolute, both surfaces glabrous, lower surface not glaucous, midrib impressed or immersed above, raised below, secondary veins 8 or 9 pairs, almost immersed above, almost immersed or slightly raised below, tertiary veins not percurrent, minor venation coarse, almost immersed above, almost immersed or slightly raised below. Vernacular name -Kenu-ong (Kenya). Notes -Beilschmiedia gynotrochioides was described by Kostermans in 1968, but was soon reduced as a synonym of B. glauciphylla by the same author (Kostermans 1970). See notes in B. glauciphylla for the differences between these two species.

Beilschmiedia gynotrochioides
Vegetatively, B. gynotrochioides approaches B. microcarpa because they share the same phyllotaxis and glabrousness. They differ from each other in venation appearance on the upper leaf surface (almost immersed in B. gynotrochioides vs raised in B. micro carpa), flower size (c. 3.5 mm diam. vs c. 2 mm diam.), fruit shape (spindle-shaped vs ellipsoid) and fruit size (c. 8 by 4 cm vs c. 3.5 by 1.5 cm). Small tree usually to 8 m tall (rarely to 18 m tall). Terminal buds ovoid, brown, pubescent with brown to reddish brown, erect, minute, curly hairs. Twigs terete, slightly angular when young, brown, pubescent with same type of hairs as on terminal buds. Petiole 1.2-1.5 cm long, flat to rounded above, drying brown, pubescent with same type of hairs as on twigs. Leaves alternate, evenly distributed, blade chartaceous to firmly chartaceous, almost glabrous, matt, elliptic, 16-26(-31) by 8-10(-13) cm, base cuneate, apex acuminate, margin flat, midrib slightly raised above, raised below, secondary veins 9-11(-13) pairs, mostly immersed above, raised below, tertiary veins weakly percurrent, minor venation relatively fine, slightly raised on both surfaces. Inflorescences axillary, paniculate, 12-45 cm long, pubescent with same type of hairs as on twigs, each bearing c. 100 or more flowers; floral pedicels of lateral divisions c. 3 mm long, pedicels of central flowers c. 4 mm long. Flowers red or pink, c. 2 mm long, c. 2.5 mm diameter. Tepals 6, equal, ovate, c. 2 by 1 mm, outside pubescent with same type of hairs as on twigs, inside pubescent with longer hairs. Stamens 9, c. 1.3 mm long, filaments in first and second whorls slightly shorter than anthers, filaments in third whorl slightly longer than anthers, all filaments pubescent with long wavy hairs, stamens in third whorl with a pair of glands; anthers glabrous, rarely pubescent only on back, obtuse at apex. Staminodes 3, sagittate. Pistil glabrous; ovary ovoid, c. 0.6 mm long; style terete, c. 0.7 mm long. Infructescences c. 15 cm long; fruiting pedicel c. 1.5 mm diam., not constricted at apex. Fruits ellipsoid, c. 4 by 1.5 cm, drying brown, surface smooth or slightly warty. Distribution -Endemic to Borneo: Sarawak (Malaysia), and C, W Kalimantan (Indonesia).

Beilschmiedia hartonoana
Habitat & Ecology -Primary mixed dipterocarp forest in lowlands or hillsides, usually on sandstone or clay soils. Altitude up to 600 m, rarely to 1200 m. Flowering: year-round; fruiting: November. Tree to 30 m tall. All parts glabrous except for the flowers. Terminal buds lanceolate, blackish. Twigs terete, slightly compressed when young, 2.5-3.5 mm diam., brown.
Petioles 1-1.7 cm long, flat to rounded above, drying almost black. Leaves opposite, evenly distributed, blade coriaceous, ovate, 6-13 by 3.5-6 cm, base rounded or obtuse, apex acute, margin flat, upper surface matt, lower surface slightly glossy, not glaucous, midrib immersed above, raised below, secondary veins 8-11 pairs, immersed above, raised below, tertiaries not percurrent, minor venation fine and immersed above, raised below. Inflorescences axillary or around terminal buds, racemose or paniculate with little branching, 2-3 cm long, almost glabrous or sparsely pubescent, each bearing less than 10 flowers; floral pedicels of lateral divisions c. Notes -Vegetatively, B. kinabaluensis approaches B. lucidula or B. glabra because they share opposite ovate leaves and glabrous terminal buds. However, B. kinabaluensis is distinct from the latter two species in having relatively smaller flowers with all the filaments shorter than anthers.
Flowers of this species are described as having only six fertile stamens here, but it is difficult to determine if this character is stable within the species. Thus far, there are only two flowering specimens collected for this species, and the flowers are young. ; Map 3 Beilschmiedia kunstleri Gamble (1910) 147. -Type: Kunstler 6854 (K), Malay Peninsula, Perak.
Distribution -Sarawak and Sabah (Malaysia), E Kalimantan (Indonesia). This species is also distributed in Thailand, the Malay Peninsula, and Sumatra.
Vernacular names -Medang (Iban); Rimapong (Murut). Note -Similarities of the floral structure suggest a close relationship between B. kunstleri and B. madang. However, they are clearly discriminated by the leaf size and shape.  . This species has also been collected from the Malay Peninsula, the Philippines, Sumatra, Java, and Bali.
Vernacular names -Lahu (Kayan); Medang merah. Note -Many specimens of B. madang from Borneo differ from the type specimen, which was collected from Java. The Bornean specimens have reddish hairs, coriaceous leaves with an enrolled margin, and coarser venation that is immersed on the upper leaf surface; the type has ochre hairs, chartaceous leaves with a flat margin, and finer venation that is slightly raised on the upper leaf surface. However, there are some intermediate specimens, which make it difficult to discriminate them as different species. More specimens, especially from Java or Sumatra, are needed to clarify the relationships. Tree to 24 m tall. Terminal buds ovoid, ochre to brown, pubescent with short, appressed, straight hairs. Twigs slightly angular, 4-5 mm diam., light brown to greenish brown, usually glabrous. Petioles 1.8-3 cm long, canaliculate above, drying greenish brown to dark brown. Leaves alternate or subopposite, evenly distributed, blade chartaceous, glabrous, elliptic, often asymmetric, 16-25(-30) by 6-12 cm; base cuneate to obtuse, apex cuspidate, margin flat, glabrous on both sides, lower surface not glaucous, midrib impressed or slightly raised above, raised below, secondary veins 8-10 pairs, immersed or slightly raised above, raised below, tertiary veins not percurrent, minor venation relatively fine, slightly raised above, slightly raised below. Inflorescences axillary, paniculate, 3-5 cm long, densely pubescent, each bearing 25-50 flowers; floral pedicels of lateral divisions c. 2 mm long, pedicels of central flowers 3-5 mm long, bracts usually persistent when flowering. Flowers yellow, c. 5 mm long, c. 6 mm diameter. Tepals 6, almost equal, ovate, c. 4 by 2 mm, pubescent outside, almost glabrous inside. Stamens 9, first and second whorls c. 2 mm long, third whorl c. 2.5 mm long, all filaments shorter than anthers, pubescent, stamens in third whorl with a pair of glands; anthers glabrous, obtuse to truncate at apex. Staminodes 3, sagittate. Pistil glabrous; ovary ovoid, c. 1 mm long; style terete, c. 1.5 mm long. Infructescences 5-15 cm long; fruit pedicels c. 2.5 mm diam., gradually thickened to 5 mm diam. below fruits, not constricted at the apex. Fruits ellipsoid, c. 2.5 by 1.5 cm, drying black, surface smooth. Distribution -In W and E Kalimantan (Indonesia), and widely scattered in Sabah and Sarawak (Malaysia). This species has also been collected from the Malay Peninsula, Sumatra, Java, and the Philippines.

Beilschmiedia maingayi
Habitat & Ecology -Primary or secondary, hill or montane forest, on yellowish or red clay soils, brownish soils, or loam soils with limestone. Altitude 10-1400 m. Flowering: January, April to July, November; fruiting: May to November. Vernacular names -Empawang Berok; Mananpuroh (Sungei); Medang. Notes -This species has often been misidentified as B. wallichiana, a poorly known species of the Malay Peninsula. Beilschmiedia maingayi, however, is distinguished from B. wallichiana by the appressed hairs on the terminal buds and the filaments shorter than anthers.
The closest species to B. maingayi in Borneo is supposedly B. oligantha, which shares similar leaf and flower shape. Beilschmiedia maingayi is distinct from B. oligantha in the indument type (appressed vs erect), flower colour (yellow vs red) and shape of the fruit pedicels (not constricted vs constricted at apex). Tree to c. 40 m tall. Terminal buds ovoid, brown, pubescent with erect, minute curly or dusty hairs. Twigs terete, slightly compressed when young, light brown to dark brown, pubescent with minute curly hairs when young. Petioles 1-2.5 cm long, rounded above, drying ochraceous to brown. Leaves alternate, evenly distributed, blade firmly chartaceous, glabrous, elliptic, 13-24 by 5-10 cm, apex acute to acuminate, base cuneate, margin flat, lower surface usually slightly glaucous, midrib almost immersed or slightly impressed above, raised below, secondary veins 8-11 pairs, immersed or slightly impressed above, raised below, tertiary veins usually not percurrent, immersed on both surfaces, minor venation fine (finest veins hardly visible on both surfaces), immersed above, almost immersed below. Habitat & Ecology -Primary or secondary forest, often near rivers, on sandy loam soils, limestone, or brown to blackish soils. Altitude up to 300 m. Flowering: from April to July, September and October; fruiting: July, October.
Notes -Large-leaved specimens of this species used to be recognized as a distinct variety, B. micrantha var. latifolia. However, the leaf size is variable in this species and is not a discriminating character.
This species approaches B. hartonoana, B. phoebeopsis, and B. tawaensis by its long inflorescences and small flowers. However, it differs from B. phoebeopsis in the leaf venation pattern and petiole appearance and from B. tawaensis in the venation pattern on the upper leaf surface. See notes on B. hartonoana for the differences between B. micrantha and B. hartonoana. Tree to 9 m tall. Terminal buds narrowly ovoid, c. 2 mm diam., blackish brown, glabrous or rarely sparsely pubescent only on the edge of buds or inside buds. Twigs terete, c. 2 mm diam., greenish brown to dark brown, glabrous. Petioles 0.7-1.5 cm long, flat above, drying brown, glabrous. Leaves opposite, evenly distributed, blade chartaceous, matt, elliptic, 11-19 by 5-8 cm, base cuneate to obtuse, margin flat, apex acuminate, both surfaces glabrous, lower surface green, midrib immersed or slightly raised above, raised below, secondary veins 7-9 pairs, slightly raised above, raised below, tertiary veins not percurrent, minor venation relatively coarse, slightly raised on both sides.
Habitat & Ecology -Primary forest on hills, on loam soils containing limestone, sandy soils, or clay soils. Altitude up to 1300 m. Flowering: late January, March; fruiting: October, February.
Note -Among opposite-leaved species of Beilschmiedia in Borneo, this species, B. glauciphylla, and B. gynotrochioides share a combination of the following characters: narrow glabrous terminal buds, elliptic leaves and short filaments. The discriminating characters between B. microcarpa and the latter two species are mainly in the fruit shape, fruit size, and flower size (for details, see notes on B. glauciphylla and B. gyno trochioides). above, drying almost black and slightly contrasting with twigs. Leaves opposite, evenly distributed, blades coriaceous, matt, narrowly elliptic, 6.5 -10 by 1.5 -3 cm, base cuneate, apex acute, margin slightly revolute, upper surface glabrous, lower surface glabrous, not known whether glaucous or not, midrib almost immersed above, slightly raised or almost immersed below, secondary veins 7-9 pairs, almost immersed above, very slightly raised below, tertiary veins not percurrent, minor venation relatively coarse, immersed above, slightly raised or almost immersed below. Inflorescences axillary, paniculate with little branching, c. 3.5 cm long, sparsely pubescent, each bearing c. 10 flowers; floral pedicels 1-2 mm long. Flowers colour unknown, c. 2.5 mm long, c. 3 mm diameter. Tepals 6, equal, ovate, c. 1.7 by 1.5 mm, sparsely pubescent on both sides. Stamens 9, filaments in first and second whorls c. 1.2 mm long, shorter than anthers, filaments in third whorl c. 1 mm long, as long as anthers, all filaments glabrous, stamens in third whorl with a pair of glands; anthers glabrous, obtuse to truncate at apex. Note -Among the opposite-leaved Beilschmiedia species of Borneo, only B. mon tanoides and B. dictyoneura have their terminal buds pubescent with appressed hairs. In addition to the terminal bud character, B. montanoides and B. dictyoneura share narrowly elliptic leaves. However, the two species differ from each other in leaf texture (coriaceous in B. montanoides vs chartaceous in B. dictyoneura) and the stamen number (9 vs 6). Tree to 10 m tall. Terminal buds ovoid, brown, pubescent with erect curly hairs. Twigs angular to sulcate, 2.5-4 mm diam., dark brown, densely pubescent with brown, erect, curly hairs. Petioles 0.5-1 cm long, flat above, drying almost black. Leaves alternate, evenly distributed, blade coriaceous, elliptic or ovate, 4-8 by 2.5-4.5 cm, base obtuse, apex obtuse to acute, margin slightly revolute, upper surface glossy, glabrous, lower surface pubescent with short, erect, curly hairs, unknown whether glaucous or not, midrib immersed above, raised below, secondary veins 6-8 pairs, slightly impressed above, raised below, tertiaries not percurrent, minor venation relatively coarse, conspicuously raised and thick above, slightly raised below. Inflorescences axillary, paniculate with little branching, c. 2.5 cm long, pubescent, each bearing c. 10 flowers; floral pedicels c. 2 mm long. Flowers colour unknown, c. 2 mm long, c. 2.5 mm diameter. Tepals 6, equal, ovate, c. 1.5 by 1 mm, both surfaces densely pubescent. Stamens 9, c. 1.1 mm long, filaments shorter than anthers, pubescent, stamens in third whorl with a pair of glands; anthers glabrous, obtuse at apex. Staminodes 3, sagittate. Pistil glabrous; ovary ovoid, c. 0.6 mm long; style terete, c. 0.6 mm long. Infructescences unknown. Distribution -Endemic to Borneo, known only from Mt Murud, Sarawak (Malaysia).
Note -This species is known only from its type specimens with only flowers, but clearly distinguished from the other Beilschmiedia species of Borneo by its alternate, small, coriaceous leaves with the lower leaf surface covered with curly hairs. Beil schmiedia madang sometimes has firmly chartaceous leaves with a pubescent lower surface and approaches this species, but differs from it in having large flowers (c. 4 mm long) with filaments longer than anthers. Tree, height unknown. Terminal buds ovoid, dark brown, pubescent with reddish brown, short, curly hairs. Twigs terete, angular when young, c. 3.5 mm diam., dark brown, almost glabrous when old. Petioles 1.8-2.2 cm long, flat above, drying dark brown.

Distribution -In W Kalimantan (Indonesia).
Habitat & Ecology -Forest or soil type unknown. Altitude around 280 m. Flowering: April.
Note -This species approaches B. madang in floral characters. The main differences between B. oblonga and B. madang are in leaf characters: B. oblonga has relatively narrow (5-6 cm) leaves with round apex and raised minor veins on the upper surface, whereas B. madang has wider (5-10 cm) leaves with an obtuse apex and immersed minor veins on the upper surface. However, some leaves of B. oblonga are almost obtuse, and many collections of B. madang outside Borneo have raised minor veins on the upper leaf surface. More specimens of B. oblonga, especially with fruits, are needed to clarify the relationship. Small tree, up to 10 m tall. Terminal buds ovoid, densely pubescent with long, erect, straight hairs. Twigs densely pubescent with long, erect, straight hairs. Petioles 1.5-2(-3) cm long, flat to slightly canaliculated above, densely pubescent. Leaves alternate, evenly distributed on branches, (14-)17-20(-24) by (4-)5-10 cm, obovate or sometimes elliptic, chartaceous, base acute, apex acuminate or cuspidate, margin flat, upper leaf surface pubescent only along midrib, lower surface sparsely pubescent with erect, long, straight hairs, lower surface glaucous, midrib almost immersed above, raised below, secondary veins 6-9(-12) pairs, almost immersed above, raised below, tertiaries not percurrent, minor venation relatively fine, slightly raised but the smallest veins immersed above, slightly raised below. Inflorescences axillary, paniculate, 1-5 cm long, each bearing a few to 30 flowers, rachis densely pubescent; floral pedicels of lateral divisions c. Habitat & Ecology -Submontane or mossy forest on sandy soils. Altitude 500-1500 m. Flowering: May; fruiting: July to October.

Beilschmiedia oligantha
Note -This species is distinguished from the other Beilschmiedia species of Borneo by its long lanceolate leaves with relatively large flowers. It was once reported as conspecific to B. wieringae (Kostermans 1969), but differs from this species in leaf shape and flower colour. Tree to 40 m tall. Terminal buds ovoid to narrowly ovoid, pubescent with dusty hairs and short, erect, curly hairs. Twigs terete, slightly angular when young, drying dark brown, pubescent with dusty hairs and short, erect, curly hairs, glabrous when old.
Vernacular names -Ambong (Labuk); Medan; Papak; Silan (Kinabatangan). Notes -This species has sometimes been confused with B. perakensis from the Malay Peninsula, presumably because they share a similar indument type, leaf shape, and flower size. Apparently, their difference was considered to be in the amount of pubescence on the lower leaf surface; a number of specimens from Borneo with less hairy leaves has been identified as B. perakensis. After comparative study of the Malaysian and Bornean specimens, however, I found that the density of pubescence on the lower leaf surface is actually a variable character in B. tawaensis, and that more reliable differences between the two species are the venation pattern, inflorescence branching pattern, and position of bracts on the flower pedicels. Beilschmiedia tawaensis is distinguished from B. perakensis by a finer venation pattern, more branched inflorescences (usually branching three times or more vs twice in B. perakensis), and bracts located at a more basal position on the pedicels. I once reported that B. tawaensis had ellipsoid fruits (Nishida 2001). However, observation of better specimens revealed that the fruit shape of B. tawaensis is instead spherical. Small tree, up to 15 m. Terminal buds narrowly ovoid, pubescent with short, erect, wavy hairs. Twigs terete, slightly angular when young, c. 1.5 mm diam., dark brown, pubescent with short, erect, curly hairs. Petioles 1-1.5 cm long, flat above, drying dark brown, sparsely pubescent. Leaves alternate, evenly distributed, blade firmly chartaceous, matt, narrowly elliptic, 10-17 by 3-5 cm, base cuneate to obtuse, apex slightly acuminate, margin flat; upper surface glabrous, lower surface pubescent with short erect hairs only along midrib, not known if glaucous or not, midrib immersed or impressed above, raised below, secondary veins 6 or 7 pairs, almost immersed above, raised below, tertiary veins not percurrent; minor venation fine, almost immersed above, slightly raised   below. Inflorescences axillary, paniculate, 4-6 cm long, pubescent with short, erect hairs, each bearing 30-60 flowers; floral pedicels of lateral divisions c. 2 mm long (c. 0.5 mm from bract), pedicels of central flowers c. 2 mm long (c. 0.5 mm from bract). Flowers yellowish green, c. 2 mm long, c. 2.5 mm diameter. Tepals 6, ovate, c. 1.5 by 1 mm, sparsely pubescent on both sides. Stamens 9, 0.6-0.9 mm long, filaments shorter than anthers in first and second whorls, as long as or longer than anthers in third whorl, filaments pubescent, stamens in third whorl with a pair of glands; anthers pubescent at apex, rarely glabrous only in third whorl, obtuse at apex. Staminodes 3, sagittate. Pistil glabrous; ovary ovoid, c. 0.7 mm long; style terete, c. 0.5 mm long. Infructescences c. 4 cm long; fruit pedicels c. 1 mm diam., constricted at apex. Fruits ellipsoid, c. 2 by 1 cm, drying dark brown, surface slightly warty.
Notes -This new species has a pubescent anther apex, which is a character usually not found in other Beilschmiedia species of Borneo. Sparsely pubescent anthers are rarely seen in a few species, such as B. hartonoana and B. phoebeopsis, but they are not pubescent on the apex, but rather on the back or only pubescent in the third whorl. In contrast, B. telupidensis has the pubescence at the top of the anthers, especially in the first and second whorls. Pubescence on the anther apex is usually a stable and reliable character for the genus, as reported for the neotropical species (Nishida 1999). Additionally, this new species can be discriminated from other Beilschmiedia species of Borneo by a combination of the following characters: erect, short, curly hairs on twigs; alternate, narrowly elliptic leaves pubescent only along the midrib on the lower surface; and immersed venation on the upper leaf surface.
The closest relative of this species in Borneo might be B. brachystachys, which shares a similar indument type on twigs (short and curly), venation appearance (veins immersed above), and flower structure (small flowers with shorter filaments in the first and second whorls). However, this new species differs from B. brachystachys in the petiole length (1-1.5 cm in B. telupidensis vs 0.5-1 cm in B. brachystachys), leaf shape (narrowly elliptic vs elliptic or obovate), leaf pubescence (pubescent only along major veins vs also on leaf tissue), pubescence on the anther apex (pubescent vs glabrous). Besides B. brachystachys, this new species resembles B. dictyoneura in the leaf appearance (narrowly elliptic, immersed veins on the upper surface) and fruit appearance (ellipsoid with warty surface). However, this new species differs from B. dictyoneura in the leaf arrangement (alternate in B. telupidensis vs opposite or subopposite in B. dictyoneura) and stamen number (9 vs 6). This species is known only by its type and one other specimen, both of which have only young fruits. These specimens share similar indument and minor leaf venation pattern with specimens of B. pilosa, but differ from B. pilosa in the obovate leaf shape, lower number of secondary veins, and non-looped secondary venation pattern. Although this species most likely belongs to Beilschmiedia, it is impossible to estimate the status or relationships to the other species without flowers. Thus far, it seems best to place this species as an imperfectly known species until more specimens become available.