A taxonomic revision of the Malesian species of Lasianthus (Rubiaceae)

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Blumea 57, 2012: 1–102 www.ingentaconnect.com/content/nhn/blumea http://dx.doi.org/10.3767/000651912X652012 RESEARCH ARTICLE INTRODUCTION Lasianthus Jack is a large genus of the Rubiaceae with more than 180 species, predominantly in the Old World. The greatest species diversity is found in tropical Asia (Robbrecht 1988), where some 160 species occur, only one extending into Australia. The species of the genus occur almost exclusively in the understory of primary forests, and have limited potential for physiological acclimation to high light levels (Cai et al. 2005). There are occasionally records from secondary or seriously disturbed forests or forest edges. Lasianthus species are usually present in large numbers in the tropical forests of Asia and may therefore represent an ecologically important element. The species of Lasianthus also show interesting distribution patterns, which are of special interest for the study of the biogeography of tropical Asia and of prevailing speciation models in tropical rainforests. The tropical Asian representatives of the genus are regarded to be extremely difficult taxonomically. The flowers and fruits are small and often shed from herbarium specimens. There are only a few diagnostic characters available for delineating the species and the majority of the characters used are quantitative features of the vegetative organs such as leaves, stipules, and bracts. It is therefore difficult to correctly determine a species by referring to old descriptions only. In most herbaria that we visited, misidentifications of Lasianthus specimens were frequent. Quite some earlier species and even newly recognized species have been described from a single type collection or a few collections only. Some types of Korthals’ species, which should be in Leiden, could not be traced, while others were destroyed in Berlin during the World War II. Even the holotype of the type species of the genus L. cyanocarpus was destroyed by a fire in 1824 (Merrill 1952). About 140 species names and numerous varieties were published based on Malesian collections (see below, Taxonomic History). All of these names except two were published before 1938. Since 1950, many more specimens from the Malesian region have been collected, but a synopsis on the genus for Malesia has not been made so far, except for two regional flora treatments, i.e. Bakhuizen van den Brink (1965) for Java (treating 29 species including 2 provisional names) and Wong (1989) for the Peninsular Malaysia (dealing with 54 species including 8 unnamed ones). Therefore, the present revision of Lasianthus from Malesia is most needed. Here we recognize 131 species from the Malesian region, of which 41 species, 3 subspecies and 3 varieties are described as new, and 3 new combinations are made for varieties. 22 species names and 15 variety names are reduced to synonyms. 10 species names and 2 varieties are treated as dubious mainly because their types could not be traced. Additionally, 11 species are excluded from Lasianthus. METHODS This paper is a classic taxonomic revision on the genus Lasianthus for Malesian species based mainly on morphological characters. The authors examined all collections from the Malesian region in AAU, BM, K, L, and U, and a selection of the Malesian collections in the herbaria BKF, KEP, MO, and SING. Type specimens in C, E, HBG, NY, P, SING, US have also been checked. The species presentation is in alphabetical order. The arrangement of synonyms is organized chronologically in homotypic paragraphs. All cited specimens are examined by the authors except a few type specimens which are indicated as ‘n.v.’. In total, all c. 3 600 specimens in K and L were examined. Additional specimens from AAU received on loan were also examined. More than 95 % of the Lasianthus collections in K and L have been identified for the present study. A small number of specimens, which do not belong to the species presently recognized, could not be treated because of imperfect state (lacking complete flowers or fruits). All published names from the Malesian region are covered in the present work. Nomenclature and typification follow the International Code of Botanical Nomenclature (McNeill et al. 2006). Lectotypifications for some of Blume’s species are made. Neotypifications are also made for some species of which we could not trace type materials and which are most probably lost. A taxonomic revision of the Malesian species of Lasianthus (Rubiaceae) H. Zhu1, M.C. Roos2, C.E. Ridsdale2 1 Xishuangbanna Tropical Botanical Garden, The Chinese Academy of Sciences, Kunming 650223, P.R. China; corresponding author e-mail: [email protected]. 2 Netherlands Centre for Biodiversity Naturalis (section NHN), Leiden University, P.O. Box 9514, 2300 RA Leiden, The Netherlands. Key words Lasiantheae Lasianthus Malesia revision Rubiaceae Abstract Based on herbarium collections, the Malesian species of the genus Lasianthus are revised. A total of 131 species including 5 subspecies and 6 varieties are recognized from the Malesian region, of which 41 species, 3 subspecies and 3 varieties are described as new, and 3 new combinations are made for varieties. 22 species names and 15 variety names are reduced to synonyms. Ten species names and 2 varieties are treated as dubious mainly because their types cannot be traced. Additionally, 11 species are further excluded from Lasianthus. All species are described and a key to Malesian Lasianthus is given. Published on 29 May 2012 2Blumea – Volume 57 / 1, 2012 TAXONOMIC HISTORY Generic delimitation Lasianthus Jack was established in 1823 based on two Sumatran species, Lasianthus cyanocarpus (later selected as the type of the genus; see below) and Lasianthus attenuatus. In the same year Blume (1823) listed several plant species under the genus name Mephitidia attributed to Reinwardt but described one species M. hexandra himself. The genus was redescribed two years later by Reinwardt (1825), but no species were mentioned by him. According to the International Code of Botanical Nomenclature (McNeill et al. 2006), the genus Mephitidia is attributed to Reinwardt ex Blume with M. hexandra Blume as type. Blume (1826–1827) realized that Mephitidia was synonymous with Lasianthus Jack and provided accounts of the Javanese species. Mephitidia continued to be used by some authors because they were aware of an earlier publication of Lasianthus Adans. (= Gordonia) for a genus in the Theaceae. De Candolle (1830) was the most significant of these authors and Korthals (1851) was the last author to use Mephitidia for the Malesian species in taxonomic publications. Later, Lasianthus Jack has been conserved for the genus treated here. Blume (1823) described from Java a monotypic genus Litosanthes, which is closely related to Lasianthus. Inherently Ridley (1923a) reduced this genus to a section of Lasianthus, however Bakhuizen van den Brink (1965) retained Litosanthes as a separate genus based on three characters: imbricate corolla, forked stipules, and pedunculate inflorescences. Deb & Gangopadhyay (1991) transferred some Indian taxa to Litosanthes, but subsequently Gangopadhyay & Chakrabarty (1992) reduced these taxa of Litosanthes to a section of Lasianthus (see section below). The circumscription of Lasianthus Jack was more or less modified by later authors, especially referring to the number of ovary locules or pyrenes. In Jack’s description Lasianthus has ovaries with 4 locules each with a single basal erect ovule, which normally develops into a drupe with 4 pyrenes. Blume (1826–1827) followed Jack’s view, but described more variation based on more Malesian species: 4–9 locules and a drupe with 4–9 pyrenes. Wight (1846) modified the genus by adding a species with a 2-locular ovary and drupes with 2 pyrenes (L. foetens Wight = Saprosma foetens (Wight) K.Schum. (1891)). Korthals (1851) established a subgenus of Dysosmia based on a Javanese species Mephitidia dichotoma Korth., which has a 2-locular ovary and a compressed bi-pyrened drupe with a thin wall. Miquel (1857) raised it to generic rank as Dysosmia Korth., and consolidated the circumscription of Lasianthus Jack based on Blume’s definition. Boerlage (1891) reduced Dysosmia Korth. to Saprosma. Baillon (1880), reputed for his wide generic concepts in general, enlarged Lasianthus Jack and moved the African genus Saldinia under Lasianthus Jack as a subgenus. Saldinia has a 2-locular ovary, which develops into a drupe with a single hard-walled pyrene. Bremekamp (1957) made a combination L. furcatus (Miq.) Bremek. based on Canthium furcatum Miq., and he sank Saprosma dichotomum (Korth.) Boerl. in L. furcatus. Dysosmia was again returned to Lasianthus by Bremekamp (1957) because its type is Saprosma dichotomum. Meanwhile, Bremekamp (1957) restored Saldinia and gave a new definition to Lasianthus as having 2–more locules in the ovary and drupes with 2–more pyrenes with a relatively soft wall (against Saldinia). Subsequently, some confusion emerged. A group of Asian species, which has 2-locular ovaries with a single basal erect ovule in each locule, and drupes with 2 pyrenes with a thin wall, as well as usually leaves with a looped venation, were treated in several different genera. Some of these species were treated as Lasianthus, some as Saprosma and some as Amaracarpus (Valeton 1911, Davis & Bridson 2004). Lasianthus as defined here has drupes or pyrenes with a thick wall. Species that show drupes with a thin wall (and do not belong to Saprosma nor Amaracarpus), are here treated as dubious taxa. Their systematic position needs further study. Piesschaert (2001) studied the carpology of Psychotrieae and gave a definition of Lasianthus as having a gynoecium with 4–12 locules that develops into a drupe with 4–12 pyrenes. In our present treatment, we restrict Lasianthus to species with drupes with 2–9 mature pyrenes with a thick wall that develop from ovaries with 2–9 locules for the Asian representatives. Petit (1964) and Denys (1981) discussed that African species have pyrenes with a pre-formed oval to circular germination lid that could easily removed. However, for Asian species, few studies of the germination lids of pyrenes have been carried out. Delimitation of the species The African taxa have to some extent been treated by Verdcourt (1976) for tropical East Africa. Denys (1981) revised the genus for Central Africa. There seems to be less work on the three American species, however, Robbrecht (1982) commented upon two of these on the occasion of a discussion of the synonymy of the Panamanian endemic genus Dressleriopsis. Important for mainland SE Asia is the work of Hooker (1880) who recognized some 52 species and placed them all in a subgeneric classification (see below, Infrageneric Classification). Subsequently regional flora accounts were produced for former ‘Indo-Chine’ (Pitard 1924) and for Thailand (Craib 1934). Later Yamazaki (1964) revised the genus for the Ryukyu Islands and Liu & Chao (1964) treated the Taiwanese species. Zhu (1994, 2002) treated the Chinese Lasianthus and then revised the genus for Thailand (Zhu 2001b). The taxa from the Indian subcontinent have received considerable attention in recent years: Deb & Gangopadhyay (1989, 1991) for India and Ridsdale (1998) for Sri Lanka. The species of the genus from the Malesian region were studied by many authors: Java (Blume (1826–1827, types in L; Bakhuizen van den Brink 1963, types in L); Java, Sumatra and, Borneo (Korthals 1851, types not found; Miquel 1857, 1861, 1869, types in U and L); Borneo (Stapf 1894, types in K, SING; Merrill 1929a, 1937, holotypes in HBG, isotypes in NY); Penin sular Malaysia (King & Gamble 1904, types mainly in SING and K, fewer in U and L; Ridley 1909, 1910, 1912, 1917a, b, 1918, 1920, 1923a, b, types in K and SING; Wong 1989); the Philippines (Elmer 1906a, b, 1908, 1911, 1912, 1913, 1934a, b, holotypes in PNH†, isotypes dispersed in A, E, K, L, MO, NY, P, U, US; Merrill 1908a, b, 1922, 1923, holotypes in PNH†, isotypes dispersed in A, K, L, MO, NY, P, U, US); New Guinea (Lauter bach 1905, holotype could be in WRSL; Schumann 1905, type in B†; Valeton 1911, 1927, type mainly in B†, fewer in L, S; Wernham 1914, 1916, 1918, types in BM). Infrageneric classification The first apparent attempt to subdivide the genus (Wight 1846) created two subdivisions ‘Bracteosae’ and ‘Nudiflorae’, without rank and not typified. These two names were taken over by Hooker (1880) and given the rank of section: Bracteatae and Nudiflorae. Hooker described two further sections ‘Stipulares’ typified by L. stipularis Blume and a non-typified section ‘Pedunculatae’. Hooker placed two of the species of Wight’s group ‘Nudiflorae’ into this section hence only one of the original species remained in ‘Nudiflorae’, i.e. L. blumeanus which is the type of the section. The type species of Lasianthus, L. cyanocarpus, is in the sect. Bracteatae and hence this is a superfluous name for sect. Lasianthus. Ridley (1923a) confused the issue further, the section including the type species also contained the species of sect. Stipulares 3 H. Zhu et al.: Revision of Malesian Lasianthus and was incorrectly called ‘Eu-Lasianthus’ and two further sections were added. These have names formerly used at generic level, ‘Mephitidia’ and ‘Litosanthes’. There was no further reference to the origin of these names in the text but it is reasonable to assume that these refer to the genera of the same name and that the typification is the same. These Javanese species were unknown in the Peninsular Malaysia and not treated directly by Ridley. Section Mephitidia, which has bracts inconspicuous or absent, is apparently the same as sect. Nudiflorae of Hooker, and sect. Litosanthes, which has forked stipules and pedunculate inflorescences, is the same as sect. Pedunculatae of Hooker. Finally, Gangopadhyay & Chakrabarty (1992) reduced the taxa included in Litosanthes back to a section of Lasianthus where Hooker’s section Pedunculatae was the oldest available name at that level. They further chose L. biermannii King ex Hook.f. as the lectotype species of this section. We accept here a division in three sections. The position in Rubiaceae In one of the classical subfamilial classifications of the Rubiaceae (Hooker 1873), Lasianthus is placed in the tribe Psychotrieae, based on the valvate aestivation of the corolla lobes and a single, basally attached ovule in each of the ovary locules. Schumann (1891) followed Hooker and retained the genus in the Psychotrieae. Petit (1964) proposed new definitions for the Psychotrieae and Morindeae, based especially on seed characters. He defined Psychotrieae as having seeds with horny endosperm and small embryos whilst the Morindeae have seeds with soft oily endosperm and large embryos. Petit therefore transferred Lasianthus to the Morindeae. Petit’s placement was followed by Verdcourt (1976) and Robbrecht (1988). However, Igersheim & Robbrecht (1993) narrowed the delimitation of the Morindeae, and Lasianthus was excluded to an uncertain position. Recent molecular work has indicated further possible relationships. Bremer (1996), working with sequences of the chloroplast gene rbcL, reported that Lasianthus forms a single clade with Pauridiantha Hook.f. with high bootstrap support values. This is very surprising in view of morphological differences as Pauridiantha has a quite different fruit, berry-like and containing a large number of seeds. Andersson & Rova (1999) sequenced the rps16 intron to investigate the phylogeny of the Rubioideae – the subfamily wherein Psychotrieae and Morindeae are placed in current systems (e.g. Robbrecht & Manen 2006). In their results Lasianthus groups together with the neotropical genus Perama Aubl. on a strongly supported branch, again in spite of morphological differences with Perama. Andersson & Rova (1999) suggested that Lasianthus should be added to the monogeneric tribe Perameae Bremek. ex Darwin, of which the position was previously uncertain as well (Robbrecht 1988). Bremer & Manen (2000), combining the results from the protein coding rbcL gene, the spacer sequence between atpB and rbcL, and Andersson & Rova’s rps16 intron data, suggested a new tribe Lasiantheae B.Bremer & Manen to include Lasianthus and Trichostachys Hook.f., the latter formerly placed in the Morindeae. Based on the chloroplast rps16 intron sequence, a phylogenetic analysis of Lasianthus was made by Xiao & Zhu (2007). Contrary to Andersson & Rova (1999), they did include Saldinia A.Rich. and Trichostachys, and in their results Lasianthus, Saldinia and Trichostachys form a strongly supported branch, with Perama as their sistergroup. In fact, Xiao & Zhu’s (2007) results are compatible with those of Andersson & Rova (1999). Piesschaert et al. (2000) added seven more genera of Rubioideae to the morphological and molecular matrix and following their cladistic analysis Lasianthus, Saldinia and Trichostachys form a closely related group and Perama is more distantly related to this group. Piesschaert et al. (2000) also found additional morphological and anatomical characters to support Petit’s (1964) ideas on the tribal characters of the Psychotrieae and Morindeae. The Psychotrieae mostly have terminal inflorescences, wood with radial multiple vessels, and seeds with small embryos and horny (not oily) endosperm. Lasianthus differs from the Psychotrieae in having axillary inflorescences paired at the nodes, solitary vessels, large embryos and soft oily endosperm. The latter seed characters are shared with the Morindeae which however, have biovulate locules and the ovules attached to the middle of the septum. Lasianthus has uniovulate locules and the ovules are attached at the base in the corner of the locule. In conclusion, the abovementioned molecular analyses brought the novel view that Lasianthus separates on a branch independent from both the Psychotrieae and Morindeae. The suggestion to recognize the group of genera Lasianthus, Saldinia and Trichostachys at the tribal level is well supported by molecular and morphological characters. Lasianthus, Saldinia and Trichostachys form a strongly supported branch in the ‘basal Rubioideae’. Further investigation is required to decide upon the position of Perama. Pending further clarification on this point it seems logical to accept the tribe Lasiantheae B.Bremer & Manen. MORPHOLOGY Life form Lasianthus species are usually shrubs, but some are understory shrubs or grow into small trees (treelets); some African and American species are creeping plants. Generally they have a monopodial axis with lateral plagiotropic shoots. Indumentum An indumentum is often present on the branchlets, petioles, the lower surface of the leaves, especially on midribs and veins, as well as on the stipules in many of the species, and if present, it is of diagnostic importance. Some species are almost glabrous, such as L. laevigatus, L. laxiflorus, and L. lucidus. A few species have a variable indumentum: young branchlets and the lower surface of leaves are hairy to subglabrous or glabrous, which has led to the separation of varieties. Stiff hairs (stiff and rough hairs) are found in L. hirsutus; hirsute hairs (rough hairs) in L. chrysotrichus, L. cyanocarpoides; villous hairs (dense, long and soft hairs) in L. curtisii, L. hirtus, L. hispidulus, L. sikkimensis and others; pubescence (short, dense and usually appressed hairs) in L. bracteolatus, L. chrysoneurus, L. chrysophyllus, L. clementis, L. euneurus, L. obliquinervis and others; very short and thin pubescence in L. chinensis, L. stercorarius; strigillose hairs (spreading rough hairs) in L. chlorocarpus, L. trichophlebus; hispid hairs (stiff, rough and spreading hairs) in L. borneensis, L. setulosus; tomentose hairs (dense, relatively short and spreading hairs) in L. chryseus, L. ferrugineus, L. tomentosus, L. undulatus and others. The species L. chrysoneurus differs from L. obscurus by having appressed-pubescent vs tomentose indumentum. Lasianthus hispidulus separates from L. clementis by only having villose vs appressed-pubescent indumentum. However, the different types of hairs are sometimes hard to distinguish, and it turns out that different concepts of hair type were used by the various authors. Here we delimit hair types as explained above. Leaf size, shape and texture The foliage varies in shape and size in the genus. Some species have relatively small leaf-blades less than 10 cm long, such as L. appressus, L. curtisii, L. lucidus. Lasianthus bifloroideus and L. biflorus have the smallest leaf-blades, less than 3 cm long. Some have leaf-blades longer than 15 cm, such as 4Blumea – Volume 57 / 1, 2012 L. chinensis and L. hirsutus. Several Bornean species have leaf-blades which are even more than 25 cm long, such as L. longibracteatus and L. megaphyllus. Most species have oblong leaf-blades, but L. linearifolius has linear leaf-blades. The leaf apex is usually cuspidate-acuminate or acute, but caudate in L. ding houanus and L. linearifolius. Most species have cuneate or acute leaf bases, but L. attenuatus, L. cyanocarpus and L. rigidus have oblique, cordate or rounded bases. Leaf texture varies from chartaceous to coriaceous. Lasianthus tenuifolius has thinly chartaceous leaf-blades; leaves of L. mucronulatus and L. oblongatus are chartaceous, while those of L. inodorus and L. vriesianus are coriaceous. The petiole varies from subsessile to several cm long. Leaf blade venation Venation and the number of secondary veins are of some diagnostic value. The veinlets (nervules) are of more diagnostic importance. Lasianthus brochidodromus, L. kinabaluensis and L. membranaceoideus have looped venation, while most Lasianthus species have open venation. Lasianthus chinensis and L. verticillatus are two easily confused species, however L. chinensis has leaves with more than 10 pairs of nerves and reticulate nervules, while L. verticillatus has leaves with less than 10 pairs of nerves and (as most species) parallel nervules. Lasianthus malaccensis and L. wightianus have conspicuously reticulate nervules. Lasianthus euneurus and L. verticillatus have parallel nervules. Lasianthus subaureus separates from L. verticillatus mainly by having reticulate nervules vs parallel nervules. However, some species have nervules between parallel and reticulate, termed subparallel and subreticulate, respectively. The venation is usually prominent beneath, but in some species prominent on both surfaces. Several species have faint and obscure nervules. Stipules The size and shape of the interpetiolar stipules are of diagnostic value within the genus. Lasianthus longifolius, L. obscurus, L. trichoplebus and L. venosus have relatively large (more than 5 mm long), triangular or lanceolate stipules. Other species have minute, inconspicuous stipules. Lasianthus stercorarius and L. vulcanicus have stipules with a swollen base. Inflorescences The inflorescences are axillary and paired at the nodes, sessile and fascicled, or pedunculate cymes. Bracts or bracteoles are present and persistent, or absent. The diagnostic importance of bracts and peduncles was emphasized by Hooker (1880). For example, L. chevalieri Pit. from mainland SE Asia is separated from L. capitatus from Indonesia mainly by the sessile inflorescence. Lasianthus laxiflorus, L. pendulus and L. robinsonii have peduncles longer than 3 cm, whereas L. brevipedunculatus and L. pergamaceus have peduncles which are only 1–3 mm long. However, length variation of the peduncles sometimes reduces its diagnostic importance. For example, L. capitatus and L. rhinocerotis have inflorescences pedunculate or subsessile. Some species, such as L. congesticymus and L. reticulatus have inflorescences extremely congested. Bracts are notably orbicular and coriaceous in L. inodorus; ovate to lanceolate in L. chrysoneurus and L. ob scurus; linear or lanceolate in L. attenuatus, L. rigidus, and L. sikkimensis; leaf-like in L. cyanocarpus and L. hirsutus, or inconspicuous or absent in L. chinensis, L. curtisii, L. lucidus, and L. tomentosus. Flowers The flowers are usually small, from several mm to a maximum of 2.5 cm long (only seen in L. longifolius). In most species they are sessile, but the flowers of L. brevipedicellatus, L. oblongus, L. pedicellatus, and L. purpureus are more or less conspicuously stalked. Some species have 4-merous flowers, such as L. floresensis, L. ridsdalei, and L. trichophlebus. Many species have 5-merous flowers. The calyx has more diagnostic characters. As it is persistent, its features can also be used for identifying material in fruiting stage. Lasianthus chrysotrichus, L. curtisii, and L. hexandrus have a calyx that is divided nearly to the base. Lasianthus lucidus, L. ridsdalei, and L. sylvestroides have calyx lobes which are conspicuously longer than the calyx tubes. Many species have calyx lobes which are nearly as long as the calyx tubes or shorter than the calyx tubes but still conspicuous. Lasianthus fordii and L. hispidulus have a calyx limb with 5 minute, triangular teeth. Lasianthus longifolius, L. subaureus, and L. verticillatus have cupular truncate calyx limbs or minute teeth at the apex. However, the length of calyx lobes is not always constant. In a few species, for example L. fordii, the calyx teeth can be variable. The corolla is easily lost when preparing specimens, and not often used for diagnostic character, because it is poorly known for many species. Imbricate corolla lobes were used as one of the characteristics for moving sect. Pedunculatae Hook.f. of Lasianthus to Litosanthes Blume by Deb & Gangopadhyay (1989, 1991). However, both imbricate and valvate corolla lobes Map 1 SE Asia showing total number of taxa for each geographical area and number of endemic taxa in brackets. 5 H. Zhu et al.: Revision of Malesian Lasianthus are observed in species of sect. Pedunculatae. The systematic significance of corolla aestivation is limited, whereas the shape of the stamens and the ovary are rarely used as diagnostic characters. However, the number of ovary locules is of importance in separating Lasianthus from closely related genera. Fruits Fruits are drupes crowned (as already mentioned above) by persistent calyx lobes. Indumentum, surface texture (smooth or warty) and morphology (ridged or not ridged and the number of ridges) of the dry fruits are more of diagnostic importance. The ridges correspond to the number of pyrenes and are especially obvious in immature seeds or seeds from herbarium material. The fruits with a thick wall and 1–9 mature pyrenes (Asian species) develop from ovaries with 3–9 locules. The surface characters of the pyrene wall and cross section are of important diagnostic value. However, in the present revision, the number of pyrenes is often used as it is relatively easy to observe, wall characters only when no other diagnostic characters are available. BIOGEOGRAPHY With regard to the geographical regions as recognized by Van Steenis (1950) and Van Welzen & Slik (2009), Borneo with 71 taxa (including subspecies and varieties), is the most biodiverse area, followed by Peninsular Malaysia with 54 taxa. Next come Sumatra with 51 taxa, Java with 43 taxa, and the Philippines with 22 taxa. In terms of endemicity, Borneo comes first again, housing 32 taxa, once more followed by Peninsular Malaysia with 18 taxa, Sumatra with 9 taxa, and the Philippines with 5 taxa (see Table 1 and Map 1). TAXONOMIC TREATMENT Lasianthus Lasianthus Jack (1823) 125, nom. cons.; Blume (1826–1827) 995; Spreng. (1830) 94; Wight (1846) 498; Miq. (1857) 314; Hook.f. (1873) 129; Kurz (1877) 30; Hiern (1877) 228; Hook.f. (1880) 178; K.Schum. (1891) 120; Prain (1903) 576; Ridl. (1923a) 149; Pit. (1924) 371; Valeton (1927) 105; Craib (1934) 207; Fl. Hainan 3 (1974) 333; T.S.Liu & J.M.Chao (1964) 118; Bakh.f. (1965) 335; K.M.Wong (1988) 367; Deb & M.Gangop. (1991) 270; H.Zhu (1994) 49; (2001b) 124; (2002) 69. — Type: Lasianthus cyanocarpus Jack (typ. cons.). Dasus Lour. (1790) 141, nom. rej. — Type: Dasus verticillata Lour. (= Lasianthus verticillatus (Lour.) Merr.). Mephitidia Reinw. ex Blume (1823) 51. — Type: Mephitidia hexandra Blume (= Lasianthus hexandrus (Blume) Blume). Litosanthes Blume (1823) 22; (1826–1827) 994. — Type: Litosanthes biflora Blume. Santia Wight & Arn. (1834) 422, nom. illeg., non Santia Savi 1799. — Type: Santia venulosa Wight & Arn. (= Lasianthus venulosus (Wight & Arn.) Wight). Subshrubs, shrubs, or rarely small trees, foetid or not. Branches and branchlets terete, sometimes compressed, rarely fistulous; lenticels inconspicuous or conspicuous. Leaves opposite, distichous, usually thinly coriaceous or chartaceous; apex acuminate, acute or cuspidate; base acute to rounded or cordate; midrib flat, depressed or slightly prominent above, usually prominent beneath; nerves generally prominent beneath, ascending at an angle of more than 45°, curved to the margin or joining the nerves above at margin; nervules parallel or reticulate. Stipules interpetiolar, usually persistent, well developed or inconspicuous, triangular, lanceolate, ovate or oblong; apex acute or obtuse. Inflorescences axillary, rarely supraaxillary, opposite, sessile, compact or cymes with peduncles, several-flowered to many-flowered; peduncles conspicuous or compressed, stout to extremely congested; bracts persistent or not, well developed or absent. Flowers small, white, sessile or pedicellate, in clusters or peduncled cymes axillary and paired at the nodes; calyx campanulate, with 3–6 teeth or lobes, rarely with cupular limb truncate at apex, persistent on fruit; corolla funnel-form or salver-shaped, several mm long to the maximum of 2.5 cm long, glabrous or hairy outside; the throat usually villous inside; the lobes 4–6, valvate or imbricate in bud; stamens 4–6, inserted on the throat of the corolla; filaments short; the anthers linear or oblong, dorsifixed, included or exserted; style linear, with 3–9 linear or lanceolate stigmas lobes; ovary 3–9-celled; ovules 1 in each cell, basal, erect. Fruit drupes small, pulpy, blue, or rarely white, black or red, usually globose, smooth or warty, rounded or ridged; pyrenes 2–9, thick walled, smooth, warty or sulcate on the abaxial face, usually triangular in transverse section. Distribution — More than 180 species in tropical areas of the world, and centred in tropical Asia. There are more than 160 species in tropical Asia, c. 20 species in Africa, 1 in Australia and 3 in tropical America. Note — Hooker’s (1880) infrageneric system has been commonly accepted although there is not an exclusive charac ter for any of the sections. Stipules, bracts and peduncles, which are key characteristics to separate sections in Hooker’s system, vary from species to species gradually in size and shape. However, some correlation among these characteristics can be seen indeed. Large stipules (large enough to cover or partly cover the inflorescence) are correlated with the sessile inflorescence and numerous bracts except for two species in which the bracts are reduced. The sect. Stipulares so defined seems a natural group. The sect. Nudiflorae, which has sessile or subsessile inflorescences and inconspicuous bracts except one species with very short peduncles, can be recognised as a loose group. The sect. Lasianthus, which has sessile cymes accompanied by conspicuous bracts, cannot be clearly separated from the sect. Pedunculatae, which has peduncled cymes, because there is a group of species with conspicuous peduncles accompanied by conspicuous bracts, such as L. moonii, L. pendulus, L. rhinocerotis. Here we therefore reduce the sect. Pedunculatae to the sect. Lasianthus. Key to the sections 1. Stipules quite large, ovate, oblong-ovate or triangular-ovate, coriaceous with membranaceous margin or membranaceous, partly or completely covering the sessile cymes .. ................................ sect. I. Stipulares Table 1 Geographical distribution of Malesian Lasianthus. Biogeographical area Number of total taxa* Number of endemic taxa* Borneo 71 32 Peninsular Malaysia 54 18 Sumatra 51 9 Java 34 1 Philippines 22 5 New Guinea 21 4 Sulawesi 20 0 Lesser Sunda Islands 11 0 Moluccas 9 0 Solomon Islands 2 1 Country Malaysia 90 34 Indonesia 81 19 Brunei 23 1 Philippines 22 5 Papua New Guinea 19 2 Singapore 9 0 Solomon Islands 2 1 * Species, subspecies and varieties together. 6Blumea – Volume 57 / 1, 2012 1. Stipules large or small, triangular, lanceolate, oblong, rarely ovate, never large enough to cover or partly cover cymes 2 2. Bracts conspicuous, linear-lanceolate, ovate, cordate or leaf-like, more that 3 mm long ....... sect. II. Lasianthus 2. Bracts inconspicuous or absent, or minute, linear, usually less than 3 mm long ..............sect. III. Nudiflorae Sect. I. Stipulares Hook.f. (1880) 179. — Type: Lasianthus stipularis Wight Key to the species 1. Stipules completely or almost covering the cymes, bracts not forming a dense and thick head if existent; leaves less than 16 cm long with 6–11 pairs of nerves ...........2 1. Stipules partly covering the cymes, bracts forming a dense and thick head; leaves usually longer than 16 cm with more than 12 pairs of nerves ................ 53. L. griffithii 2. Bracts numerous, longer than 5 mm ................ 3 2. Bracts few, less than 5 mm long ................... 4 3. Bracts linear to narrowly lanceolate, up to 1.5 cm long, villous ............................. 117. L. stipularis 3. Bracts ovate-triangular to lanceolate, 7–8 mm long, glabrous except hairy margins ...................52. L. glaber 4. Branchlets and leaves glabrous; leaves oblong; calyx with a shallowly cupular limb, truncate or minute-teethed at apex .......................... 100. L. pseudo-stipularis 4. Branchlets hirsute; leaves ovate-elliptic with hirsute nerves beneath; calyx without a shallowly cupular limb but with lanceolate lobes with barbate hairs at margin 9. L. barbiger Sect. II. Lasianthus – sect. Bracteatae Hook.f. (1880) 179, nom. illeg. — Type: Lasianthus cyanocarpus Jack. – sect. Pedunculatae Hook.f. (1880) 190. — Type: Lasianthus biermannii King ex Hook.f. (lecto, designated by Gangopadhyay & Chakrabarty 1992). Key to the species 1. Cymes sessile, subsessile or shortly tuberculate ...... 2 1. Cymes conspicuously pedunculate, peduncles more than 1 mm long ...................................31 2. Bracts longer than 5 mm .........................3 2. Bracts shorter than 5 mm ....................... 17 3. Leaves with subcordate or rounded, more or less oblique bases ........................................4 3. Leaves with cuneate or acute, equal-sided bases......9 4. Leaf nerves conspicuously depressed above, 10–13 pairs; drupes glabrous .................. 33. L. crassinervis 4. Leaf nerves flat or slightly prominent above, usually less than 10 pairs; drupes hairy or glabrous ..............5 5. Plants glabrous; leaves with a faint intramarginal vein; bracts coriaceous, oblong-lanceolate to oblong, obtuse at the apex; calyx lobes oblong to oblong-ovate, obtuse at the apex 30. L. cordatus 5. At least stipules and bracts hirsute; leaves without an intramarginal vein; bracts chartaceous, not oblong-lanceolate or oblong, acuminate or acute at the apex; calyx lobes triangular or lanceolate, acute or acuminate at the apex ........6 6. Branches glabrous, enlarged (swollen) at nodes; leaves hirsute on midrib and nerves beneath, obovate to obovateoblong, coriaceous; drupes glabrous......106. L. rigidus 6. Branches usually densely tomentose or villous, not enlarged (swollen) at nodes; leaves usually densely tomentose, villous or setaceous beneath, usually oblong, membranaceous or chartaceous; drupes hairy .......................7 7. Branchlets densely spreading villous; bracts paired, with a long linear apex ............... 113. L. sogerensis 7. Branchlets densely tomentose; hairs not spreading; bracts not paired, but numerous, acuminate ..............8 8. Bracts cordate to broadly lanceolate, somewhat leaf-like ..............................37. L. cyanocarpus 8. Bracts linear to lanceolate, not leaf-like..6. L. attenuatus 9. Branchlets and leaves usually stiffly rusty-hirsute; bracts large, cordate, orbicular, ovate or ovate-lanceolate, leaflike, more than 1 cm broad ............56. L. hirsutus 9. Branchlets and leaves not stiffly rusty-hirsute; bracts relatively small, oblong-lanceolate, lanceolate, linear or subulate, less than 0.5 cm broad ................10 10. Bracts oblong-lanceolate or lanceolate, with a conspicuous midrib...................................... 11 10. Bracts linear or linear-lanceolate, without a conspicuous midrib......................................12 11. Branchlets thinly appressed-pubescent; leaves narrowly oblong, less than 3 cm broad, thinly appressed-pubescent on midrib and nerves beneath; bracts 2 in each axil, oblong-lanceolate, less than 2 mm broad, glabrous above, appressed-pubescent hairs on midrib and margin beneath .......................... 86. L. oblongatus 11. Branchlets densely brown-hirsute; leaves oblong-elliptic, more than 3 cm broad, densely hirsute beneath; bracts numerous, lanceolate, more than 2 mm broad, hirsute .. ........................... 36. L. cyanocarpoides 12. Stipules ovate-triangular, up to 7 mm long ........... ................................. 40. L. ellipticus 12. Stipules not ovate-triangular, relatively small, less than 4 mm long or inconspicuous .................... 13 13. Bracts linear-lanceolate; drupes glabrous ............ ..............................112. L. sikkimensis 13. Bracts filiform or subulate-linear; drupes hairy ...... 14 14. Bracts filiform, more than 8 mm long, forming a dense head .................................. 34. L. crinitus 14. Bracts subulate-linear, not forming a dense head .... 15 15. Leaves oblong to oblong-lanceolate, less than 3 cm broad ................................ 32. L. coronatus 15. Leaves elliptic-oblong or obovate-elliptic, more than 3 cm broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16. Branchlets spreading tomentose; leaves elliptic, spreading tomentose beneath ...............69. L. ledermannii 16. Branchlets subglabrous or appressed-pubescent; leaves obovate-elliptic, appressed-pubescent or subglabrous beneath ......................... 13. L. bracteolatus 17. Leaves very large, 22–30 by 7–12 cm; drupes verrucose, with 6 conspicuous ridges . . . . . . . . . 76. L. megaphyllus 17. Leaves less than 20 cm long, less than 6 cm broad; drupes without 6 conspicuous ridges . . . . . . . . . . . . . . . . . . . 18 18. Leaves thickly coriaceous; bracts 2, spatulate-obovate, 3 by 2 mm; flowers 4-merous; drupes 3–4............. ..........................88. L. obovatibracteatus 18. Leaves coriaceous or chartaceous; bracts more than 2, not spatulate-obovate; flowers 4–6-merous; drupes 4–5 . 19 19. Leaves coriaceous; bracts orbicular or ovate-orbicular 20 19. Leaves chartaceous, membranaceous or subcoriaceous; bracts neither orbicular nor ovate-orbicular .........21 20. Leaves small, ovate-lanceolate, 8–11 by 1.5 cm with deeply depressed midrib and nerves above; calyx lobes ovate, longer than the tube ...........38. L. depressineurus 7 H. Zhu et al.: Revision of Malesian Lasianthus 20. Leaves relatively large, more than 10 cm long and more than 2.5 cm broad; midrib and nerves not depressed above; calyx lobes ovate-lanceolate, not longer than the tube .. ................................. 60. L. inodorus 21. Branchlets and leaves glabrous; stipules and bracts glabrous except barbate margins ..........8. L. barbatus 21. Branchlets and leaves hairy beneath; stipules and bracts hairy.......................................22 22. Stipules more than 5 mm long; bracts ovate or ovate-triangular..................................... 23 22. Stipules less than 5 mm long; bracts linear to linear-lanceolate or lanceolate ..........................25 23. Leaves with acute bases; nerves 3–6 pairs; petioles 1.5–2 cm long; bracts small, less than 2 mm long; calyx lobes triangular, 1.5 mm long, glabrous internal surface; drupes conspicuously 5–8-ridged ......... 85. L. obliquinervis 23. Leaves with cuneate to obtuse bases; nerves 8–10 pairs; petioles less than 1 cm long; bracts longer than 2 mm; calyx lobes lanceolate, longer than 1.5 mm, hirsute both surfaces; drupes without conspicuous ridges .......24 24. Densely spreading tomentose hairs on branches, leaves beneath, stipules, bracts, calyx and drupes . . . . . . . . . . .................................89. L. obscurus 24. Thinly appressed-pubescent hairs on branches, leaves beneath, stipules, bracts and calyx; drupes subglabrous ............................. 23. L. chrysoneurus 25. Stipules triangular; branchlets and leaves beneath appressed-pubescent or puberulous; nervules parallel, bracts lanceolate to subulate; drupes with 4 pyrenes . . . . . . 26 25. Stipules lanceolate, linear, subulate or inconspicuous; branchlets and leaves beneath hirsute or tomentose; nervules parallel or reticulate; bracts linear, linear-lanceolate or linear-subulate; drupes with 4 or more pyrenes ...27 26. Branchlets, leaf venation beneath, stipules and bracts appressed-puberulous; drupes glabrous except persistent calyx lobes .................. 50. L. foxworthyanus 26. Branchlets, leaf venation beneath, stipules and bracts very densely appressed-pubescent; drupes puberulous..... ................................... 80. L. mollis 27. Leaf base cordate or slightly cordate and more or less oblique; nerves more than 12 pairs; calyx lobes up to 5 mm long; corolla up to 1.5 cm long ...... 97. L. pilosus 27. Leaf base cuneate, acute or subrounded, not oblique; nerves less than 10 pairs; calyx lobes less than 5 mm long; corolla less than 1.2 cm long ....................28 28. Leaves more than 13 cm long; bracts numerous; fruits verrucose .............................. 104. L. ridleyi 28. Leaves less than 12 cm long; bracts not numerous; drupes smooth ....................................29 29. Leaves subrounded at base; stipules inconspicuous; bracts linear-lanceolate; pyrenes 8 . . . . . . . . . . 4. L. appressus 29. Leaves cuneate or acute at base; stipules linear-subulate or narrowly lanceolate; bracts linear; pyrenes 5–6 ... 30 30. Leaves lanceolate; leaf nervules strongly reticulate; calyx lobes lanceolate, c. 2.5 mm long .........98. L. politus 30. Leaves oblong or elliptic oblong; leaf nervules parallel; calyx lobes linear, 3–4 mm long ...122. L. sylvestroides 31. Peduncles usually more than 4 cm long ........... 32 31. Peduncles usually less than 4 cm long ............ 33 32. Leaves oblong-ovate to ovate; nerves 4–6 pairs; petioles c. 5 mm long; stipules triangular, less than 2 mm long .. ........................... 46. L. filipedunculatus 32. Leaves oblong; nerves 8–9 pairs; petioles 2–3 mm; stipules triangular lanceolate, 4–5 mm long ....107. L. robinsonii 33. Peduncles very slender, less than 0.3 mm diam .....34 33. Peduncles relatively robust, more than 0.3 mm diam . 35 34. Branchlets appressed hirsute; leaves elliptic-obovate, less than 2.5 cm long; leaf bases cuneate; stipules very small, inconspicuous; bracts small, subulate or inconspicuous; ovary 4-locular; drupes ovoid with 4 pyrenes ......... .................................. 11. L. biflorus 34. Branchlets long spreading villose; leaves oblong-lanceolate, more than 5 cm long; leaf bases rounded to subcordate; stipules linear, c. 4 mm long; bracts linear to ovate, leaf-like, 6–10 mm long; ovary 8-locular; drupes globose with 6–8 pyrenes .................. 45. L. filiformis 35. Plant totally glabrous ..........................36 35. Branchlets and leaf nerves beneath hairy or more or less appressed-pubescent .........................38 36. Main peduncles up to 3 cm long, trifurcate; stipules up to 5 mm long; calyx c. 3 mm long, with c. 1 mm long subulate teeth ............................68. L. laxiflorus 36. Main peduncles less than 1.5 cm long, bifurcate; stipules less than 4 mm long; calyx 8–10 mm long, with 5–6 mm long oblong-lanceolate or lanceolate lobes .........37 37. Leaves with looped venation; nerves 4–5 pairs; stipules very small, subulate, less than 1 mm long; drupes with 4 pyrenes; pyrenes very warty ...... 51. L. furcatoides 37. Leaves without looped venation; nerves 8–10 pairs; stipules triangular, 3 mm long; drupes with 6 pyrenes; pyrenes smooth .........................64. L. laevigatus 38. Peduncles 2–3.5 cm long ......................39 38. Peduncles less than 2 cm long .................. 41 39. Branchlets, leaves beneath and peduncles sparsely hirsute; leaves chartaceous with acute base; nerves 5–6 pairs; petioles 6–8 mm long; bracts linear; flowers 4-merous, pediceled ........................65. L. lancilobus 39. Branchlets, leaves beneath and peduncles densely spreading villose; leaves coriaceous with subcordate, rounded or obtuse bases; nerves 7–10 pairs; petioles shorter than 6 mm; bracts linear or filiform; flowers 5-merous, sessile ...40 40. Leaves ovate-lanceolate, with an acuminate apex and a subcordate base; midrib and nerves deeply depressed above; nervules subparallel; petioles c. 1 mm long; bracts numerous, filiform, up to 1.5 cm long; corolla glabrous .. .................................93. L. pendulus 40. Leaves ovate-oblong with a caudate apex and a rounded or obtuse base; nerves flat above, nervules parallel; petioles 5 mm long; bracts 2, linear, 7–8 mm long; corolla densely villous ..........................75. L. malaiense 41. Bracts 2, ovate, orbicular or cordate, leaf-like, more than 1 cm broad .................................42 41. Bracts more than 2, linear or subulate, not leaf-like, less than 5 mm broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 42. Branchlets and leaves densely spreading yellowish hirsute; leaves oblong, with slightly asymmetric base, 10–17 by 3–5 cm; peduncles 7–8 mm long, bracts relatively small, c. 1.5 by 1 cm, with shortly cuspidate apex; calyx lobes 5, 2–3 mm long; pyrenes 5 ............18. L. cailinianus 42. Branchlets and leaf nerves, bracts and calyx densely appressed-pubescent; leaves slightly obovate-oblong, 26–34 by 6–8 cm; peduncles c. 2–3 mm long; bracts large, c. 2 by 2 cm, with a c. 5 mm long cuspidate apex; calyx lobes 4, 1.5 mm long; pyrenes 4 ........ 24. L. chrysophyllus 43. Leaves 26–46 cm long; stipules c. 1 cm long; peduncles very short, 1–2 mm long; bracts up to 3 cm long ...... ............................71. L. longibracteatus 43. Leaves less than 16 cm long; stipules less than 5 mm long; peduncles short or long; bracts less than 1.5 cm long 44 8Blumea – Volume 57 / 1, 2012 44. Bracts linear, 6–15 mm long, hirsute; cymes capitate 45 44. Bracts linear or subulate, less than 5 mm long; hirsute or pubescent; cymes not capitate ..................46 45. Leaf bases subrounded; nerves 9–12 pairs; bracts numerous, usually forming a dense head; peduncles usually very short; calyx lobes linear-lanceolate, c. 3 mm long; drupes with 5 pyrenes . . . . . . . . . . . . . . . . . . 103. L. rhinocerotis 45. Leaf bases cuneate or acute; nerves 7–8 pairs; bracts not forming a dense head; peduncles up to 1.5 cm long; calyx lobes subulate-linear, up to 12 mm long; drupes with 6 pyrenes ..........................19. L. capitatus 46. Branchlets, leaves beneath and peduncles densely tomentose .......................................47 46. Branchlets, leaves beneath and peduncles appressed-pubescent or glabrous ...........................48 47. Peduncles 1–3 mm long; calyx lobes oblong to lanceolateoblong, 3 mm long ...........15. L. brevipedunculatus 47. Peduncles longer than 5 mm; calyx lobes linear, 5–6 mm long ........................ 121. L. sumatraensis 48. Leaves linear-lanceolate, less than 2 cm broad, long-caudate at the apex; calyx lobes triangular, c. 0.8 mm long . ..............................39. L. dinghouanus 48. Leaves not linear-lanceolate, more than 2.5 cm broad, acuminate or cuspidate at the apex; calyx lobes ovateoblong or lanceolate, 1.5–4 mm long ............. 49 49. Branchlets, leaves beneath and peduncles densely appres sed-pubescent; nerves 3–6 pairs; cymes trifurcate; flowers with 3–5 mm long pedicels, 4-merous; calyx lobes lanceolate, c. 1.5 mm long; drupes with 4 conspicuous ridges; pyrenes 4 ............... 67. L. laxifloroideus 49. Branchlets, leaves beneath and peduncles thinly pubescent or glabrous; nerves 7–8 pairs; cymes bifurcate; flowers sessile, 5-merous; calyx lobes oblong, 3–4 mm long; drupes with 5 ridges; pyrenes 5 ......... 110. L. scalariformis Sect. III. Nudiflorae Hook.f. (1880) 184. — Type: Lasianthus blumianus Wight Key to the species 1. Cymes very shortly peduncled; branchlets conspicuously sulcate; leaves large, coriaceous, 18–25 by 5–8 cm; calyx obconic-cupular, truncate or minutely 4-toothed ....... ............................. 95. L. pergamaceus 1. Cymes sessile or tuberculate; branchlets not sulcate; leaves various; calyx various ....................2 2. Leaves less than 3 cm long .........10. L. bifloroideus 2. Leaves more than 3 cm long .....................3 3. Leaves linear, 10–14 by 0.9–1.8 cm, long-caudate at the apex, glabrous ...................70. L. linearifolius 3. Leaves otherwise..............................4 4. Leaves with conspicuously looped venation; branchlets and leaves glabrous except sometimes puberulous nerves beneath .....................................5 4. Leaves without looped venation; branchlets and leaves glabrous or hairy ..............................7 5. Leaves lanceolate, nerves 9–12 pairs; drupes puberulous and slightly warty ............. 16. L. brochidodromus 5. Leaves oblong or oblong-lanceolate; nerves less than 9 pairs; drupes glabrous and conspicuously warty...... 6 6. Leaves subrounded at the base; nerves 6–8 pairs; drupes with conspicuous 8 ridges; pyrenes with 2 furrows on the abaxial face . . . . . . . . . . . . . . 77. L. membranaceoideus 6. Leaves acute to cuneate at the base; nerves 3–4 pairs; drupes with 3–4 obtuse ridges; pyrenes without furrow on the abaxial face . . . . . . . . . . . . . . . . 63. L. kinabaluensis 7. Flowers on pedicels 2 mm long or more ............ 8 7. Flowers sessile or subsessile pedicels less than 1 mm long .......................................13 8. Leaves thinly membranaceous; nervules weak, inconspicuous; stipules broadly ovate ..120. L. submembranifolius 8. Leaves chartaceous to thickly coriaceous; nervules conspicuous; stipules otherwise .....................9 9. Leaves elliptic, oblong-elliptic or oblong, large, 4.5–8 cm broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9. Leaves lanceolate, ovate-lanceolate, oblong-lanceolate, less than 4 cm broad .......................... 11 10. Leaves chartaceous or subcoriaceous with acute bases ...........................14. L. brevipedicellatus 10. Leaves thickly coriaceous with subrounded bases ..... ...............................129. L. vriesianus 11. Leaves; nervules reticulate; pedicels 8–10 mm long; drupes verrucose, c. 6 mm diam, with 4 conspicuous obtuse ridges; pyrenes 4.......................101. L. purpureus 11. Leaves; nervules parallel; pedicels 2–3 mm long; drupes 3–4 mm diam, with 4–5 (smooth) or 7–8 (verrucose) obtuse ridges; pyrenes 4–5 or 7–8 .................12 12. Stipules less than 1 mm long; calyx with triangular teeth; drupes smooth, with 4–5 obtuse ridges; pyrenes 4–5 .. .............................. 92. L. pedicellatus 12. Stipules 3–4 mm long; calyx truncate; drupes verrucose, with 7–8 obtuse ridges; pyrenes 7–8 ...87. L. oblongus 13. Leaves narrowly oblong-lanceolate, narrowly lanceolate or linear-lanceolate, 5–6 times longer than broad, longer than 12 cm .................................14 13. Leaves otherwise ............................17 14. Plants densely tomentose; leaves 23–25 by 3–4 cm; nerves 16–18 pairs; calyx with 7–8 lanceolate lobes ......... .................................41. L. eriocalyx 14. Branchlets glabrous or pubescent; leaves 9–19 by 1.5–3 cm; nerves less than 15 pairs; calyx with 4–5 teeth or with an almost truncate limb . . . . . . . . . . . . . . . . . . . . . . . . 15 15. Nerves 7 pairs; stipules narrowly triangular, c. 4 mm long; calyx almost truncate or minutely 5-toothed .......... ............................1. L. acuminatissimus 15. Nerves 8–14 pairs; stipules triangular, 2–3 mm long; calyx with 4–5 conspicuous teeth ....................16 16. Branchlets glabrous; leaves with densely hirsute margin; nerves 8–11; petioles 12–15 mm long .............. ............................ 57. L. hirtimarginatus 16. Branchlets densely appressed-pubescent; leaves without hirsute margin; nerves 12–14 pairs; petioles 4–5 mm long ...............................3. L. angustifolius 17. Drupes usually with 2 pyrenes (only in Borneo) ..... 18 17. Drupes with 3 or more pyrenes . . . . . . . . . . . . . . . . . . 20 18. Leaves membranaceous; drupes ovoid, very rugose, without cork-like structure between pyrenes; pyrenes rugose on the abaxial face . . . . . . . . . . . 78. L. membranaceus 18. Leaves coriaceous; drupes ellipsoid, smooth, with cork-like structure between pyrenes; pyrenes smooth on the abaxial face .......................................19 19. Plants glabrous; leaves with subrounded base, nervules conspicuously prominent on both surfaces; petioles 5–7 mm long; stipules c. 2 mm long; calyx tube 3 mm long, not expanded at limb ................. 84. L. nigrescens 19. Branchlets, midrib and nerves beneath and stipules puberulous; leaves with cuneate base; nervules depressed 9 H. Zhu et al.: Revision of Malesian Lasianthus or flat above, prominent beneath; petioles 1–1.5 cm long; stipules 4–6 mm long; calyx tube c. 1 mm long, limb c. 1–1.5 mm long, more or less expanded............ ..............................109. L. sabahensis 20. Plants spreading stiffly hirsute; leaves with subrounded and oblique base; nerves 10–12 pairs; stipules narrowly triangular or lanceolate, 4–5 mm long ... 2. L. acutatus 20. Plant otherwise ..............................21 21. Stipules ovate or ovate-triangular or triangular with a swollen base and a membranaceous margin, 3–6 mm long 22 21. Stipules otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 22. Branchlets, leaves, stipules and calyx densely spreading villose; calyx with triangular lobes; drupes villose ....... ................................... 58. L. hirtus 22. Branchlets, leaf midrib and nerves beneath (on lower surface), stipules and calyx thinly puberulous or glabrous; calyx subtruncate, with 4–5 minute teeth at the apex; drupes subglabrous or glabrous .................23 23. Branchlets, leaf midrib and nerves beneath, stipules and calyx thinly puberulous; leaves with acuminate-caudate apex; nervules obscure above, subparallel; calyx urceolate; drupes with 6–9 pyrenes .........115. L. stercorarius 23. Branchlets (except the youngest part), leaf midrib and nerves beneath and calyx quite glabrous; leaves with caudate apex; nervules distinct above, parallel; calyx shortly cupular; drupes with 4–5 pyrenes ...130. L. vulcanicus 24. Stipules longer than 6 mm......................25 24. Stipules shorter than 6 mm ..................... 29 25. Stipules broadly triangular or oblong-ovate, or oblong-lanceolate, coriaceous, glabrous or pubescent, persistent; drupes with sharp ridges . . . . . . . . . . . . . . . . . . . . . . . 26 25. Stipules lanceolate, triangular-lanceolate, linear-lanceolate or subulate, chartaceous, hirsute or strigose, often deciduous; drupes without sharp ridges ................27 26. Stipules appressed-pubescent; leaves 12–16 by 3–4.5 cm; nerves 9–12 pairs; nervules subparallel; petioles 2–3 mm long; calyx lobes triangular-lanceolate, 1–1.5 mm long; drupes 8–10-ridged; pyrenes 4–5 .... 127. L. venosus 26. Stipules glabrous; leaves 20–33 by 6–11 cm; nerves 13–18 pairs; nervules reticulate; petioles 5–10 mm long; calyx cupular with minutely teeth at the apex; drupes 6-ridged; pyrenes 6........................72. L. longifolius 27. Nervules parallel; calyx lobes 4; ovary 4-locular ....... ............................ 125. L. trichophlebus 27. Nervules reticulate to subreticulate; calyx lobes 5; ovary 5-locular ...................................28 28. Nerves 11 pairs; stipules triangular-lanceolate, 6–7 mm long, hirsute ................... 90. L. palawanensis 28. Nerves 6–8 pairs; stipules subulate, 8 mm long, appressedpubescent ....................... 91. L. papuanus 29. Stipules 4–5 mm .............................30 29. Stipules shorter than 4 mm ..................... 32 30. Leaves very large, 20–32 by 6–10 cm; nerves 10–12 pairs; calyx 7–8 mm long, lobes oblong, 4–5 mm long ...... .......................... 99. L. pseudolongifolius 30. Leaves less than 20 cm long; nerves 6–7 pairs; calyx shorter than 5 mm, with minute teeth or lobes triangularsubulate ...................................31 31. Branchlets densely appressed-pubescent; nervules parallel; calyx lobes 5–7, triangular-subulate, as long as the tube; pyrenes 5–7 ............. 114. L. solomonensis 31. Branchlets sparsely spreading hirsute; nervules subreticulate; calyx limb minutely toothed at the apex; pyrenes 4 ..............................21. L. chlorocarpus 32. Flowers 4-merous; calyx constricted at the base; drupes ellipsoid or ovoid, very rugose, narrowed at the base; pyrenes 1–4, deeply furrowed on the abaxial face ... 33 32. Flowers 4–5-merous; calyx not constricted at base; drupes globose or subglobose, verrucose or smooth, not narrowed at the base; pyrenes 4 or more, not deeply furrowed on the abaxial face . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 33. Branchlets and leaves almost glabrous; leaves chartaceous to subcoriaceous; cymes conspicuously tuberculate (short congested peduncles, up to 5 mm long); calyx with 4 triangular lobes; the lobes c. 1 mm long; corolla tube slender, more than 5 mm long .......... 27. L. congesticymus 33. Branchlets and leaf nerves beneath more or less appressed puberulous, pubescent or strigose; leaves coriaceous; cymes sessile or shortly tuberculate; calyx truncate or minutely toothed or with minutely triangular lobes; corolla tube stout, less than 3 mm long .................34 34. Branchlets thinly puberulous or sparsely strigose; leaves 8–13 by 3–4 cm; nervules 3–6 pairs; petioles 4–5 mm long; cymes sessile; calyx truncate or minutely toothed . ............................... 29. L. constrictus 34. Branchlets densely appressed-pubescent; leaves 19–26 by 6–9 cm; nerves 6–7 pairs; petioles 5–10; cymes shortly tuberculate; calyx with 4 minutely triangular lobes ..... ...............................102. L. reticulatus 35. Calyx with a cupular limb, truncate or minutely toothed at the apex ...................................36 35. Calyx with conspicuously lobes ................. 39 36. Leaves 18–30 cm long ..............7. L. baasianus 36. Leaves less than 18 cm long .................... 37 37. Plant glabrous; leaves chartaceous; petioles c. 2.5–3.5 mm long; drupes without an enlarged cupular calyx . . . . ...............................28. L. conspicuus 37. Branchlets and leaf venation beneath pubescent or puberulous; leaves coriaceous; petioles longer that 5 mm, drupes with an enlarged cupular calyx ............38 38. Leaves with conspicuously reticulate nervules; nerves 10– 12 pairs; petioles 10–15 mm long; drupes 5–6-ridged .. ...............................118. L. subaureus 38. Leaves with parallel nervules; nerves 5–9; petioles 5–10 mm long; drupes without ridges .... 128. L. verticillatus 39. Calyx divided to the base, or calyx lobes conspicuously longer than the calyx tube . . . . . . . . . . . . . . . . . . . . . . 40 39. Calyx not cleft to the base, calyx lobes shorter than the calyx tube or just as long as the calyx tube ........48 40. Calyx lobes oblong, obtuse at the apex 12. L. borneensis 40. Calyx lobes lanceolate or lanceolate-triangular, acuminate or acute at the apex ..........................41 41. Leaves large, more than 12 cm long .... 66. L. latifolius 41. Leaves relatively small, usually less than 12 cm long ...........................................42 42. Leaves glabrous or glabrous except appressed-hirsute or sparsely hirsute midrib and nerves and/or margin beneath; branchlets glabrous or sparsely hirsute or more or less appressed-hirsute; calyx lobes narrowly lanceolate with a long-acuminate apex or oblong; drupes glabrous or sparsely hirsute..................................43 42. Leaves very hairy beneath; branchlets very hairy but not appressed-hirsute; calyx lobes broadly lanceolate, usually acute at the apex; drupes hairy ..................45 43. Leaves ovate or ovate-elliptic, usually caudate at the apex; nervules parallel; flowers 5-merous; calyx lobes narrowly lanceolate with a long acuminate apex; drupes smooth, without ridges; pyrenes 5 ............. 73. L. lucidus 16 Blumea – Volume 57 / 1, 2012 bracts. Lasianthus attenuatus has linear to lanceolate bracts, while L. cyanocarpus has larger and cordate, ovate to broadly lanceolate, somewhere leaf-like bracts. The syntype of L. densifolius in L (Junghuhn s.n.) agrees exactly with the type of L. wallichii. However, Hooker’s description and the cited specimen Wall. Cat. 8441 (K) as L. densifolius does not agree with Miquel’s type. It represents a different variety. King & Gamble (1904), Ridley (1923a) and Wong (1989) followed Hooker’s conception and mixed them. Here we treat the different variety as a new variety of L. attenuatus. b. var. minor H.Zhu, var. nov. A var. attenuato foliis minoribus lineario-oblongis 2 cm latis, stipulis triangulelanceolatis 5–8 mm longis, bracteis et floribus minoribus differt. — Typus: J.F.Maxwell 82-72 (holo L), Singapore, Nee Soon, swamp forest. Lasianthus densifolius Miq. (1857) 321, p.p. quoad syntype: Horsfield s.n. (K), Blambangan, Java. Lasianthus densifolius auct. non Miq.: Hook.f. (1880) 182, cited specimen Lasianthus? Wall. Cat. 8441 in part; King & Gamble (1904) 122; Ridl. (1923a) 156; K.M.Wong (1989) 368, p.p.]. This variety has relatively small linear-oblong leaves which are less than 2 cm broad, triangular-lanceolate and hirsute stipules which are 5–8 mm long, relatively small and subulate to linear bracts which are 5 mm long, and relatively small flowers which are 8 mm long. Distribution — Peninsular Malaysia, Singapore, Indonesia (Sumatra). Habitat & Ecology — In swamp forest and lowland forests. Flowering and fruiting between January and March. Note — The var. attenuatus differs from var. minor in having oblong leaves which are more than 2 cm broad, inconspicuous stipules, relatively large bracts and flowers. 7. Lasianthus baasianus H.Zhu, sp. nov. — Fig. 2 Proximus L. longifolio Wight, sed ramulis, foliis subtus et calycibus puberulis, stipulis minoribus, floribus minoribus atque limbis calycis ad maturitatem fructus 2 mm longis differt. A L. pseudolongifolio H.Zhu ramulis foliisque subtus puberulis et limbis calycis cupulatis truncatis differt. — Typus: Nooteboom 4189 (holo L), Borneo, Bukit Raya, 112°45'E, 0°45'S, 1300 m alt., in primary dipterocarp forest, 5 Dec. 1982. Treelet, 3 m high; branchlets robust, compressed or subangular, fistulous, c. 5 mm diam, pubescent to glabrescent. Leaves: blades elliptic-oblong, 18–30 by 6–11 cm, coriaceous, glabrous above, puberulous beneath, apex cuspidate, base acute to subrounded, midrib prominent above, conspicuously prominent beneath, puberulous beneath, nerves depressed to flat above, prominent beneath, 9 pairs, ascending at an angle of 50–70°, curved to the margin, puberulous beneath, nervules clear above, prominent beneath, subreticulate, puberulous beneath; petioles 1–3 cm long, puberulous. Stipules triangular, 3 by 3 mm, pubescent. Inflorescences sessile, or shortly tuberculate; bracts inconspicuous. Flowers fascicular, sessile; calyx 3–4 mm long, pubescent, limb cupular, truncate or 4 minutely teeth; corolla 6–7 mm long, upper part puberulous external surface, lobes 4, lanceolate, 3 mm long, puberulous external surface, flocculous internal surface. Drupes subglobose, 5 by 6 mm, with 5 conspicuous ridges, puberulous, crowned by persistent cupular calyx limbs; calyx limbs 2 mm long; pyrenes 5, smooth on the abaxial face, subelliptic in transverse section. Distribution — Endemic to Borneo. Habitat & Ecology — In lowland primary dipterocarp forest between 100–500 m altitude. Flowering between April and December; fruiting in January. Additional specimens examined. Sarawak, Paul Chai S.34053 (L); Central Kalimantan, J.P. Mogea et al. 4019 (L); Borneo, Bukit Raya, Veldkamp 8003 (L), Nooteboom 4735 (L); Central Kalimantan, A.C. Church et al. 888 (L), 937 (K, L). Note — This species is close to L. longifolius Wight. It differs from the latter by having pubescent branchlets, leaves beneath and calyx; smaller stipules (3 by 3 mm) and relatively small flowers (up to 8 mm long); drupe crowned by 2 mm long calyx limb. While L. longifolius has glabrous branchlets, leaves and calyx; stipules up to 1 cm long; flowers up to 3 cm long; drupes crowned by 5 mm long enlarged cupular calyx limb. It differs from L. pseudolongifolius by having pubescent branchlets, leaves beneath, and cupular, truncate calyx limbs. 8. Lasianthus barbatus H.Zhu, sp. nov. — Fig. 3 Proximus L. glabro Ridl., sed foliis lanceolatis, 1.3–2.5 cm latis, nervulis subtus conspicuis, parallelis atque stipulis et bracteis minoribus differt. — Typus: T. Suppiah s.n. KEP 104817 (holo L), Peninsular Malaysia, Kelantan, Ulu Sat FR., in primary forest, 13 June 1968. Shrub, 2 mm high; branchlets terete, c. 2 mm diam, glabrous, brown. Leaves: blades lanceolate, 8–12 by 1.3–2.5 cm, subcoriaceous, glabrous except sparsely hirsute nerves beneath, apex caudate, aristate, arista 2 mm long, base acuminate to acute, midrib slightly depressed above, prominent beneath, glabrous, nerves flat above, prominent beneath, 5–6 pairs, ascending at an angle of 50–60°, curved to margin, sparsely hirsute beneath, nervules obscure above, conspicuous beneath, parallel, glabrous; petioles c. 5 mm long, glabrous. Stipules ovate-triangular to broadly triangular, 4–5 by 4 mm, glabrous except barbate margin. Inflorescences axillary, sessile; bracts ovate-triangular, glabrous except barbate margin, 4–5 by 2–4 mm; bracteoles ovate-triangular, glabrous except barbate margin. Flowers fascicular, sessile. Drupes subglobose, 4–5 by 4 mm, with 5 obtuse ridges, glabrous; calyx lobes persistent, lanceolate, 1 mm long, hirsute; pyrenes 4–5, smooth on the abaxial face, sulcate on the adaxial and the side faces in transverse section. Distribution — Endemic to Peninsular Malaysia. Habitat & Ecology — In primary forest. Fruiting in June. Note — This species has stipules and bracts with barbate margins as L. glaber. It differs from the latter by having lanceolate leaves less than 2.5 cm broad, nervules conspicuous beneath, parallel, relatively small stipules and bracts; while L. glaber has elliptic-oblong leaves, 12–18 by 4–5.5 cm, nervules obscure on both surfaces, relatively large stipules and bracts. It is close to L. foxworthyanus, but differs from the latter by having glabrous branchlets and leaves, leaves lanceolate, nerves 5–6, stipules and bracts ovate-triangular, glabrous except barbate margins, drupes with 4–5 pyrenes. 9. Lasianthus barbiger Ridl. Lasianthus barbiger Ridl. (1917b) 52. — Type: Robinson & Kloss s.n. (holo BM; iso K), Sumatra, Korinchi. Shrubs, up to 3 m high; branchlets usually compressed, 2–3 mm diam, hirsute. Leaves: blades ovate-elliptic, 9–13 by 3.5–5 cm, subcoriaceous, sparsely hirsute on midrib and nerves both sides, apex shortly caudate or cuspidate, base cuneate, midrib and nerves slightly depressed above, prominent beneath, nerves 10–11 pairs, ascending at an angle of c. 70°, curved to the margin, nerves prominent and sparsely hirsute beneath, subreticulate; petioles c. 1 cm long, hirsute. Stipules 0.5–1 cm long, lanceolate-oblong to ovate-oblong, membranaceous, with a broad base and an obtuse apex, sparsely hirsute. Cymes sessile; bracts inconspicuous. Flowers sessile, 4-merous; calyx tubular, with a campanulate limb, calyx tube c. 2 mm long, glabrous, calyx limb with 4 lanceolate c. 1.5 mm long lobes, barbate at margin; corollas unknown. Drupes ovoid, 5 by 8 mm, 17 H. Zhu et al.: Revision of Malesian Lasianthus Fig. 3 Lasianthus barbatus H.Zhu. a. Habit; b. detail of lower surface of leaf; c. stipule; d. bract; e. fruit; f. cross section of fruit (all: T. Suppiah s.n. KEP 104817, holo L). 3 mm 2 cm 2 mm 2 mm 2 mm 18 Blumea – Volume 57 / 1, 2012 Fig. 4 Lasianthus bifloroideus H.Zhu. a. Habit; b. detail of lower surface of leaf showing parallel nervules; c. calyx; d. fruit; e. cross section of fruit (all: Paul Chai S.33809, holo L). 1 mm 2 cm 2.5 mm 19 H. Zhu et al.: Revision of Malesian Lasianthus narrow at base, with 4 obtuse ridges, crowned by persistent calyx lobes, glabrous; pyrenes 4, verrucose on the abaxial face. Distribution — Endemic to Sumatra. Note — This species has large membranaceous stipules similar to those of the closely related species L. pseudostipularis and L. stipularis. It differs from L. pseudo-stipularis by having hirsute branchlets, ovate-elliptic leaves with hirsute nerves beneath, calyx lobes lanceolate, barbate at margin. It differs from L. stipularis by having hirsute branchlets, ovateelliptic leaves with hirsute nerves beneath, inconspicuous bracts, calyx lobes lanceolate, barbate at margin. 10. Lasianthus bifloroideus H.Zhu, sp. nov. — Fig. 4 Similis L. bifloro (Blume) M.Gangop. & Chakrab., sed ramulis rare puberulis, foliis ovato-lanceolatis glabris, cymis sessilis vel subsessilis, lobis calycis ovato-oblongis longioribus tubo et pyrenis conspicue verrucatis differt. — Typus: Paul Chai S.33809 (holo L; iso K), Borneo, Lubok Antu, Bukut Peninjau, in valley bottom, 13 Mar. 1974. Small shrub, 1–2 m high, branchlets terete, slender, less than 1 mm diam, puberulous. Leaves: blades very small, ovatelanceolate, 1.5–3 by 0.5–1 cm, chartaceous, glabrous on both surfaces, apex acuminate, mucronate, base acute; midrib slightly prominent above, prominent beneath, nerves slender, flat above, prominent beneath, 5 pairs, ascending at an angle of c. 60°, curved to margin, nervules weak, slender, parallel; petioles short, less than 1 mm long, puberulous. Stipules lanceolate, c. 1 mm long, puberulous. Inflorescences sessile or subsessile, bracts absent. Flowers very small; calyx c. 1 mm long, sparsely puberulous, calyx lobes 4, ovate-oblong, c. 0.7 mm long; corolla not seen. Drupes subglobose, 4–5 mm diam, glabrous, 4-ridged, crowned by 4 c. 1 mm long ovate-oblong calyx lobes; pyrenes 4, warty on abaxial face, orbicular in transverse section. Distribution — Endemic to Borneo (Lukok Antu). Habitat & Ecology — Valley bottom near river. Flowering and fruiting in March. Note — This species is closely related to L. biflorus, but differs from the latter by having branchlets thinly puberulous, leaves ovate-lanceolate glabrous, cymes sessile or subsessile, calyx lobes ovate-oblong longer than calyx tube, drupes more warty. Lasianthus biflorus has branchlets hirsute, leaves subrhombic to elliptic-obovate, hirsute midrib and nerves beneath, cymes with 5–15 mm long peduncles, drupes less warty. 11. Lasianthus biflorus (Blume) M.Gangop. & Chakrab. — Map 3 Lasianthus biflorus (Blume) M.Gangop. & Chakrab. (1992) 338; H.Zhu (2002) 100. — Litosanthes biflora Blume (1826–1827) 994. — Type: Blume s.n. (lecto L, designated by Zhu 2002, barcode L0000722), Java. Lasianthus gracilis King & Gamble (1904) 132; Ridl. (1923a) 169; K.M.Wong (1989) 367. — Type: Ridley 2904 (lecto SING, designated by Wong 1988), Peninsular Malaysia. Lasianthus gainii M.Gangop. & Chakrab. (1992) 338. — Litosanthes gracilis Hayata (1912) 113. — Syntypes: Hayata & Sasaki s.n. (n.v.), Kusukusu; G. Nakahara s.n. (n.v.), Botanrosha. Shrubs, 1–2 m high; branchlets terete, densely hirsute. Leaves: blades chartaceous, subrhomboid or elliptic-obovate, 1.5–2.5 by 0.8–1.5 cm, apex acute, often mucronulate, base cuneate, glabrous above, hirsute beneath, midrib depressed above, prominent beneath, hirsute beneath, nerves flat above, prominent beneath, 6–8 pairs, hirsute beneath, nervules obscure on both surfaces; petioles 1–2 mm long, hirsute. Stipules small, inconspicuous. Cymes with peduncles 0.5–1.5 cm long, hirsute; bracts small, inconspicuous. Flowers with pedicels c. 1 mm long; calyx campanulate, c. 2 mm long, glabrous, calyx limb with 5 triangular teeth; corolla 2–4 mm long, glabrous outside, lobes ovate. Drupes ovoid, c. 4–5 mm diam, glabrous, verrucose, 4-grooved; pyrenes 4. Distribution — China (S Yunnan, Hainan, Taiwan), mainland SE Asia, Malaysia (Peninsular, Borneo), Philippines, Indonesia (Sumatra, Java, Kalimantan, Sulawesi, Lesser Sunda Islands). 12. Lasianthus borneensis Merr. Lasianthus borneensis Merr. (1929a) 292. — Type: Elmer 21212 (iso CAS, HUH, K, L, MO, SING, U), Elmer 21656 (para MO, SING), Borneo, Tawao. Lasianthus sepalinus Ridl. (1934) 120, syn. nov. — Type: Ridley 9703 (holo K), Borneo, Bongaya, Labuk Bay; Clemens 9467 (para SING), Sandakan. Shrubs, c. 1–2 m high; branchlets terete, c. 2 mm diam, densely appressed-pubescent, virid to brown. Leaves: blades oblong to oblong-lanceolate, 10–22 by 3–5.5 cm, coriaceous to subcoriaceous, glabrous and nitid above except midrib, densely shorthirsute or hispid beneath, apex shortly caudate-acuminate, base acute to subrounded, slightly asymmetric, midrib, nerves and nervules flat to slightly prominent above, conspicuously prominent beneath, nerves 8–10 pairs, ascending at an angle of 60–70°, curved to the margin; nervules parallel; petioles 8–10 mm long, densely pubescent. Stipules oblong-ovate to lanceolate, 2–3 mm long, densely pubescent. Cymes sessile; bracts absent. Flowers sessile, 5-merous; calyx campanulate, Map 3 Distribution of Lasianthus biflorus (Blume) M.Gangop. & Chakrab. (▲) and L. obscurus (Blume ex DC.) Miq. (¡). 20 Blumea – Volume 57 / 1, 2012 Fig. 5 Lasianthus brevipedicellatus H.Zhu. a. Habit; b. lower surface of leaf showing subparallel nervules; c. fruit; d. cross section of fruit (all: Y.P. Ching S.55288, holo L). 5 mm 2 mm 5 mm 21 H. Zhu et al.: Revision of Malesian Lasianthus hirsute, calyx tube c. 2 mm long, lobes 5, oblong, c. 3 mm long; corolla tube cylindrical, c. 6 mm long, pubescent outside, villose inside; corolla lobes 5, oblong, acute at apex, 3 mm long, pubescent outside, floccose inside; anthers oblong, c. 1.5 mm long. Drupes globose, c. 5 mm diam, with 5 obtuse ridges, hirsute, crowned by 5 large oblong calyx lobes; pyrenes 5. Distribution — Malaysia (Borneo) and Brunei. Note — This species is easily recognized by its hispid leaves, relatively large calyx and oblong calyx lobes with an obtuse apex. It is closely related to L. setulosus also from Borneo. Lasianthus setulosus differs in having densely spreading long hispid on branchlets and both surfaces of leaves, a relatively small calyx, 3 by 3 mm, with ovate-triangular calyx lobes with acute apex. 13. Lasianthus bracteolatus Miq. Lasianthus bracteolatus Miq. (1869) 246; Bakh.f. (1965) 337. — Type: Junghuhn s.n. (holo L, barcode L 0057503; iso L, barcode L0057504, L, barcode L0057505), Sumatra, Tapanuli. Shrubs; branchlets terete, 2–3 mm diam, densely appressedpubescent to subglabrous, brown. Leaves: blades elliptic to obovate-elliptic, 9–13 by 4–6 cm, subcoriaceous, glabrous above, pubescent on midrib, nerves and nervules beneath, apex shortly caudate to cuspidate, base rounded to subrounded, slightly asymmetric, midrib and nerves flat above, prominent beneath, nerves 7–8 pairs, ascending at an angle of 60–70°, curved to the margin, nervules obscure above, pro minent beneath, subparallel; petioles 4–5 mm long, densely pubescent. Stipules triangular-subulate, c. 4 mm long, densely pubescent. Cymes sessile; bracts linear, 5–6 mm long, densely pubescent. Flowers sessile; calyx campanulate, densely pubescent, tube c. 2 mm long, lobes 5, subulate, c. 1 mm long; corolla not seen. Drupes globose, c. 3 mm diam, pubescent, crowned by subulate calyx lobes; pyrenes 4–5, smooth on the abaxial face. Distribution — Indonesia (Java and Moluccas). Note — This species has elliptic to obovate-elliptic leaves with 7–8 pairs of nerves, bracts linear, and drupes with 4–5 pyrenes. It differs from L. rigidus by having relatively small leaves with rounded base, longer petioles, bracts linear and shorter, calyx lobes subulate, small and smaller pubescent drupes. 14. Lasianthus brevipedicellatus H.Zhu, sp. nov. — Fig. 5 Similis L. vrieseano Miq, sed foliis chartaceis vel subcoriaceis, basi acutis differt. A L. perakensis King & Gamble foliis glabris et drupis pedicellatis 5–6-porcatis apice truncatis differt. — Typus: Ching S.55288 (holo L), Borneo, Sarawak, Sg. Palog, 850 m alt., in dipterocarp forest, 24 Sept. 1987. Treelet, 3 m high; branchlets terete, compressed in the youngest part, c. 3 mm diam, puberulous, fistulous. Leaves: blades elliptic to oblong-elliptic, 19–23 by 5–8 cm, chartaceous to subcoriaceous, glabrous on both surfaces, apex cuspidate to falcate-acuminate, mucronate, base acute, midrib slightly depressed or flat above, conspicuously prominent beneath, nerves prominent on both surfaces, 7–9 pairs, ascending at an angle of 50–60°, curved to margin, glabrous, nervules prominent on both surfaces, subparallel, glabrous, margin recurved; petioles 1–1.5 cm long, puberulous or subglabrous. Stipules triangular, 1.5–2 mm long, often deciduous. Inflorescences sessile; bracts absent. Flowers not seen. Drupes stalked, fascicular; pedicels short, 1.5–2 mm long, slender, glabrous; drupes subglobose, sparsely puberulous, warty, 5–6 conspicuously obtuse ridges; pyrenes 5–6, slightly warty on the abaxial face. Distribution — Malaysia (Peninsular and Sarawak). Habitat & Ecology — In montane forest between 850–1000 m altitude. Fruiting in September. Additional specimen examined. Peninsular Malaysia, Gunong Tapis, 3000 ft, KEP-FRI 10973 (L). Note — This species is characterized by its stalked drupes, very small stipules, bracts absent and glabrous leaves with subparallel nervules prominent on both surfaces. It is very closely related to L. vriesianus, but differs from the latter by having chartaceous to subcoriaceous leaves with acute base and smaller flowers. Lasianthus vriesianus has thickly coriaceous leaves with subrounded base. In L. vriesianus, only the type with few immature drupes has been seen. More other diagnostic characteristics between these two species cannot be recognized because of insufficient collections. This species is also similar to L. perakensis, but differs from the latter by having glabrous leaves, pediceled drupes with 5–6 ridges, while L. perakensis has puberulous leaves on nerves beneath, sessile drupes with 10-ridged pyrenes and crowned by persistent obtuse spreading calyx lobes. 15. Lasianthus brevipedunculatus H.Zhu, sp. nov. — Fig. 6 Proximus L. nervosus King & Gamble, sed foliis majoribus 4.5–8 cm longis 2–3 cm latis, cymis brevipedunculatis, floribus 5-meris, lobis calycis triangulatooblongis et pyrenis 5 differt. — Typus: Nagamasu 3359 (holo L), W Sumatra, Gunung Sago, 2100–2260 m alt., in upper montane forest around the summit, 30 Oct. 1988. Shrub, c. 2 m high; branchlets terete, c. 1.5 mm diam, densely brown-yellowish tomentose. Leaves elliptic-oblong to ovateoblong, 4.5–8 by 2–3 cm, coriaceous, glabrous above, puberulous beneath, densely tomentose hairs on midrib and nerves beneath, apex acute or cuspidate, base subrounded, midrib depressed above, prominent beneath, nerves flat or slightly depressed above, prominent beneath, 5–6 pairs, ascending at an angle of 50–60°, curved to margin, nervules prominent on both surfaces, parallel, margin hirsute; petioles 4–6 mm long, densely tomentose. Stipules triangular, 2 mm long, densely tomentose. Inflorescences shortly peduncled; peduncles 1–3 mm long, densely tomentose; bracts 1–2, linear, c. 2 mm long, tomentose. Flowers sessile; calyx hirsute external surface, lobes 5, oblong or lanceolate-oblong, 3 mm long, tube very short, less than 1 mm long; corolla 4–5 mm long, tomentose external surface, lobes 5, triangular-oblong, 2–3 mm long; stamens 5; ovary with 5 locules. Drupes c. 4 mm diam, subglobose, subglabrous, crowned by 5 calyx lobes; pyrenes 5. Distribution — Indonesia (W Sumatra). Habitat & Ecology — In upper montane forest at 2100 m altitude. Note — This species is closely related to L. nervosus from Malaysian mountains. It differs from the latter by having relatively large leaves (4.5–8 by 2–3 cm), cymes shortly peduncled, flowers 5-merous, calyx lobes triangular-oblong, and drupes with 5 pyrenes. Lasianthus nervosus has leaves of 2.5–3.5 by 1.5 cm, cymes sessile, flowers 4-merous, calyx with triangular teeth, and drupes with 4 pyrenes. 16. Lasianthus brochidodromus H.Zhu, sp. nov. — Fig. 7 Proximus L. membranaceoideus H.Zhu, sed foliis lanceolatis 2–2.5 cm latis, drupis minoribus puberulis obtuse 4–5-porcatis et pyrenis abaxiali leviter verrucosis differt. — Typus: Gaerlan et al. PPI 23712 (holo L), Philippines, Luzon, Quezon, Alabat Isl., 175 m alt., in dipterocarp forest, 25 March 1996. Shrub, c. 1.5 m high; branchlets terete, compressed at nodes, c. 1.5 mm diam, glabrous. Leaves: blades lanceolate, glabrous except puberulous nerves and margin beneath, 10–15 by 2–2.5 cm; subcoriaceous, apex acuminate, margin slightly undulate and recurved, base acute, midrib slightly prominent above, strongly prominent beneath, nerves prominent on both surfaces, 9–12 pairs, ascending at an angle of c. 90°, curved near margin 22 Blumea – Volume 57 / 1, 2012 Fig. 6 Lasianthus brevipedunculatus H.Zhu. a. Habit; b. detail of lower surface of leaf showing indumentum and parallel nervules; c. detail of node showing stipule and axillary inflorescences; d. corolla split opened; e. calyx split opened; f. fruit (all: H. Nagamasu 3359, holo L). 2 cm 2 mm 2 mm 2 mm 23 H. Zhu et al.: Revision of Malesian Lasianthus Fig. 7 Lasianthus brochidodromus H.Zhu. a. Habit with fruits; b. detail of leaf; c. detail of node showing axillary fruits; d. cross section of fruit (all: F. Gaerlan et al. PPI23712, holo L). 1 cm 2 cm 5 mm 1 cm 24 Blumea – Volume 57 / 1, 2012 Fig. 8 Lasianthus bruneensis H.Zhu. a. Habit with inflorescences; b. lower surface of leaf showing reticulate nervules; c. detail of node showing stipule and axillary inflorescences; d. fruit (all: G.T. Prance et al. 30620, holo L). 1 cm 2 cm 2.5 mm 25 H. Zhu et al.: Revision of Malesian Lasianthus and joining the nerves above, forming an intramarginal vein (looped venation), internerved vein weak, ascending from the midrib at an angle of c. 90°, combination to the main nerves, nervules conspicuous on both surfaces, reticulate; petioles 1–1.5 cm long, dark, glabrous or sparsely pubescent. Stipules small, subulate, c. 1.5 mm long, pubescent. Inflorescences sessile; bracts absent. Flowers not seen. Drupes in fascicles, sessile, globose, c. 5 mm diam, puberulous, slightly warty, with 4–5 obtuse ridges, crowned by very small calyx teeth; pyrenes 4–5, slightly warty on the abaxial face, orbicular in transverse section. Distribution — Endemic in Alabat Isl., Quezon, Philippines. Habitat & Ecology — In dipterocarp forest. Fruiting in March. Note — This species is characterized by its lanceolate leaves with a looped venation. It seems to be close to L. membranaceoideus, but differs from the latter by having leaves lanceolate, 2–2.5 cm broad, drupes puberulous, c. 5 mm diam, with 4–5 obtuse ridges, pyrenes slightly warty on the abaxial face, orbicular in transverse section. Lasianthus membranaceoideus has leaves oblong, 3–4 cm broad, drupes glabrous, c. 7 mm diam, conspicuously 8-ridged, pyrenes sulcate on the abaxial face, triangular in transverse section. 17. Lasianthus bruneensis H.Zhu, sp. nov. — Fig. 8 Proximus L. subaureo Craib, sed foliis basi rotundatis conspicue asymmetricis, stipulis minoribus 1 mm longis atque calycibus dentatis differt. — Typus: Prance et al. 30620 (holo L; iso K), Brunei, Bukit Belalong, 909 m alt., in lower montane forest around summit, 23 Feb. 1992. Treelet, 2 m high; branchlets terete, puberulous, glabrescent or glabrous. Leaves: blades linear-lanceolate to oblong-lanceolate, 8–14 by 1.5–3 cm, coriaceous, glabrous or sparsely puberulous on leaf nerves beneath, apex attenuate to acuminate or caudate, margin recurved, base rounded, asymmetric, midrib conspicuously prominent on both surfaces, nerves flat above, prominent beneath, 6–7 pairs, ascending at an angle of 75–85°, curved to the margin, nervules conspicuous beneath, reticulate; petioles nigrescent, 1 cm long, sparsely puberulous. Stipules very small, triangular, 1 mm long, puberulous. Inflorescences sessile or short tuberculate, bracts inconspicuous. Flowers in fascicles sessile; calyx campanulate, 2 by 2 mm, puberulous, limb with 5 subulate-triangular 0.5 mm long teeth; corolla not seen. Drupes c. 4 mm diam, subglobose, subglabrous, black, crowned by persistent small subulate-triangular calyx teeth; pyrenes 4–5, slightly rugose on the abaxial face, triangular in transverse section, with thickened wall. Distribution — Brunei. Habitat & Ecology — In lower montane forest between 800–1160 m altitude. In sandstone soil. Flowering and fruiting between October and February. Additional specimens examined. Brunei, Bukit Tudal, Davis A.P. 481. Note — This species is close to L. subaureus. It differs from the latter by having leaves with rounded and asymmetric bases, very small stipules, calyx not cupular, calyx limb with subulatetriangular teeth. Lasianthus subaureus has leaves with acute to acuminate base, stipules 3 mm long, calyx cupular, with truncate limb. 18. Lasianthus cailinianus H.Zhu, sp. nov. — Fig. 9 Proximus L. chrysophyllo Miq., sed plantis dense patenter fulvi-hirsutis, foliis minoribus oblongis basi leviter asymmetricis, pedunculis cymae robustis 7–8 mm longis, bracteis minoribus apice brevicuspidatis, lobis calycis 5, 2–3 mm longis et pyrenis 5 differt. Similis L. cyanocarpo Jack, sed plantis dense fulvi-hirsutis, foliis majoribus 10–17 cm longis basi abrupte acutis vel subrotundatis, nervis 14-binatis et pedunculis cymae robustis 7–8 mm longis differt. A L. hirsuto (Roxb.) Merr. plantis dense fulvi-hirsutis, foliis basi abrupte acutis vel subrotundatis, nervis 14-binatis, stipulis minoribus 3 mm longis et cymis pedunculatis differt. — Typus: Ismael 3/21.8.1957 (holo L), W Sumatra, east of Pajakumbuh, 500–1000 m alt., in primary forest, 21 Aug. 1957. Shrub, c. 2.5 m high, branchlets terete, or slightly compressed in the youngest part, robust, c. 3–4 mm diam, densely spreading yellowish hirsute. Leaves: blades oblong, 10–17 by 3–5 cm, thickly chartaceous, stiffly spreading yellowish hirsute on both surfaces, apex acute, base abruptly acute or subrounded, slightly asymmetric, midrib densely stiffly hirsute above, strongly prominent and densely hirsute beneath, nerves depressed above, strongly prominent beneath, densely hirsute, c. 14 pairs, ascending at an angle of c. 80°, curved gradually to margin, nervules obscure above, conspicuous beneath, subreticulate, hirsute, margin slightly recurved, densely hirsute; petioles 5–7 mm long, densely hirsute. Stipules lanceolate-triangular, c. 3 mm long, densely hirsute. Inflorescences pedunculate; peduncles 7–8 mm long, robust, densely yellowish hirsute; bracts 2, leaf-like, ovate, c. 1.5 by 1 cm, with shortly cuspidate apex and rounded or subcordate base, and with midrib and nerves like the leaves, densely yellowish hirsute on both surfaces; bracteoles numerous, outer ones like bracts but smaller, interior ones ovate to linear, c. 1 cm long, densely hirsute; calyx c. 4–5 mm long, densely hirsute, lobes 5, lanceolate, 2–3 mm long, densely hirsute on both surfaces; corolla densely hirsute. Drupes c. 4 mm diam, subglobose, densely hirsute, crowned by persistent linear-lanceolate, c. 5 mm long, hirsute calyx lobes; pyrenes 5. Distribution — Indonesia (W Sumatra). Habitat & Ecology — In montane primary forest between 500–1000 m altitude. Notes — This species is characterized by its densely spreading yellowish hirsute hairs, peduncled cymes with leaf-like bracts and bracteoles. It is closely related to L. chrysophyllus, but differs from the latter by having branchlets, leaves both surfaces densely spreading yellowish hirsute; leaves relatively small, oblong, with slightly asymmetric base, 10–17 by 3–5 cm; peduncles 7–8 mm long, bracts relatively small with shortly cuspidate apex; flowers 5-merous; calyx lobes 5, 2–3 mm long. Lasianthus chrysophyllus has branchlets and leaf venation, bracts and calyx densely appressed-pubescent; leaves larger, slightly obovate-oblong, 26–34 by 6–8 cm; peduncles c. 2–3 mm long; bracts larger, with a c. 5 mm long cuspidate apex; flowers 4-merous; calyx lobes 4, c. 1.5 mm long. It is also related to L. cyanocarpus, but differs from the latter by having plants covered by densely yellowish hirsute, leaves relatively big (10–17 cm long) with abruptly acute to subrounded base and 14 pairs of nerves, cymes with 7–8 mm long robust peduncles. While L. cyanocarpus has much soft pilose hairs on branchlets, leaves and bracts; leaves relatively small (8–13 cm long) with rounded to cordate oblique base and less than 10 pairs of nerves; cymes sessile or subsessile. This species is also similar to L. hirsutus, but differs from the latter by having densely yellowish hirsute hairs, leaves with 14 pairs of nerves and abruptly acute or subrounded base, stipules 3 mm long, cymes peduncled. Lasianthus hirsutus has rusty hirsute hairs, leaves with 7–10 pairs of nerves and acute to cuneate base, stipules 8–10 mm long, and cymes sessile. 19. Lasianthus capitatus Blume Lasianthus capitatus Blume (1826–1827) 996; Miq. (1857) 320; (1869) 246; Merr. (1921) 577; Bakh.f. (1965) 337. — Mephitidia capitata (Blume) DC. (1830) 453. — Type: Blume s.n. p.p. (the part from Java, non Sumatra) (lecto L, designated here, barcode L0310036), Java. Shrubs; branchlets terete, c. 3 mm diam, densely tomentose. Leaves: blades oblong-elliptic to oblong-lanceolate, 10–15 by 3–3.5 cm, coriaceous, glabrous above, tomentose beneath, apex acuminate, base acute or cuneate, midrib and nerves flat 32 Blumea – Volume 57 / 1, 2012 Fig. 10 Lasianthus coriaceifolius H.Zhu. a. Habit with inflorescences; b. lower surface of leaf showing reticulate nervules; c. detail of node showing stipule and axillary inflorescences; d. fruit; e. cross section of fruit (all: B. Lee S.52556, holo L). 4 mm 2 cm 2 mm 5 mm 33 H. Zhu et al.: Revision of Malesian Lasianthus 34. Lasianthus crinitus Hook.f. Lasianthus crinitus Hook.f. (1880) 182. — Type: Wall. Cat. 8441, p.p. (holo K), Peninsular Malaysia, Penang, or Singapore. Shrubs; branchlets terete, c. 3 mm diam, densely brown-tomentose. Leaves: blades oblong, 10–15 by 2.5–4 cm, coriaceous, glabrous above, densely brown-tomentose beneath, apex acute to acuminate, base rounded, margin reflexed, midrib and nerves flat to slightly depressed above, prominent beneath, nerves 8–9 pairs, ascending at an angle of c. 50°, curved to the margin, nervules distinct above, distinct and prominent beneath, parallel; petioles 3–4 mm long, densely brown-tomentose. Stipules inconspicuous. Cymes sessile; bracts numerous, filiform, up to 1 cm long, densely tomentose, forming a dense head. Flowers clustered, sessile; calyx and corolla not seen. Drupes globose, c. 3 mm diam, densely brown-tomentose, 5-ridged, crowned by 5 linear calyx lobes which are c. 2 mm long and tomentose; pyrenes 5. Distribution — Peninsular Malaysia. Note — This species has branchlets and leaf blades beneath rusty-tomentose, cymes sessile, bracts filiform, dense and numerous. The type is a fragment mounted with two other taxa, i.e. L. attenuatus and L. attenuatus var. minor on the same sheet. Only one specimen from Peninsular Malaysia (FRI 15716 from Fraser’s hill of Pahang) in L that we examined matches the type and description. This species is closely related to L. lineolatus Craib by its indumentum and bracts. 35. Lasianthus curtisii King & Gamble — Map 4 Lasianthus curtisii King & Gamble (1904) 128; Ridl. (1923a) 161; Craib (1934) 209; T.Yamaz. (1964) 352, p.p.; T.S.Liu & J.M.Chao (1964) 126, f. 5; H.Zhu (1994) 64; (1998) 151; H.S.Lo (1999) 92; H.Zhu (2001b) 126; (2002) 86. — Lectotype: Curtis 284 (lecto SING, designated by Zhu 2002; isolecto K), Peninsular Malaysia, Penang. Lasianthus formosensis Matsum. var. hirsuta Matsum. (1901) 17. — Type: Owatari s.n. (holo TI, n.v.), Taiwan. Lasianthus kwangtungensis Merr. (1929b) 323. — Type: Tsiang 55 (holo SCBI; iso K), China, Hongkong. Lasianthus condorensis Pierre ex Pit. (1924) 385. — Syntypes: Balansa 2658 (P), Vietnam Tonkin, Bavi; Eberhardt 1981 (P), Annam, Quang-tri; Harmand 916 (P), Cochinchina [Vietnam], Condor. Lasianthus pseudo-lucidus King & Gamble (1904) 128; Ridl. (1923a) 168. — Type: Wray 4109 (holo SING), Peninsular Malaysia, Perak, 5000 ft. Shrubs, 1–2 m tall; branches and branchlets terete, c. 1.5–2 mm diam, densely villous. Leaves: blades ovate-lanceolate or elliptic-lanceolate, 5–9 by 2–3 cm, subcoriaceous, glabrous above, densely and softy villous beneath, apex cuspidateacuminate, base cuneate or obtuse, midrib and nerves flat above, prominent beneath, nerves 4–6 pairs, ascending at an angle of 50–60°, curved to the margin, nervules obscure above, distinct beneath, subparallel; petioles 4–5 mm long, villose. Stipules minute, inconspicuous. Cymes sessile; bracts absent. Flowers sessile; calyx divided nearly to the base into 5 lanceolate up to 5 mm long, acuminate lobes, hirsute outside; corolla 7–8 mm long, 5-lobed, outside hispid, inside villous, the throat hairy. Drupes ovoid-globose, hirsute, 4–5 mm diam, crowned by the long persistent calyx lobes; pyrenes 5. Distribution — China, Japan (Amami, Kyushu, Ryukyu), Vietnam, Thailand, Peninsular Malaysia, Indonesia (Sumatra). Note — Wong (1989) reduced L. pseudo-lucidus to L. curtisii. Lasianthus pseudo-lucidus is very closely related to L. curtisii in having calyx almost the same as the latter. However, it has narrower leaves, which is a little different from the typical L. curtisii. 36. Lasianthus cyanocarpoides Valeton Lasianthus cyanocarpoides Valeton (1927) 108; Merr. & L.M.Perry (1946) 219. — Type: Schlechter 17805 (holo S; iso K, L), Papua New Guinea, Kaiser-Wilhemsland, 1000 m, 29 May 1908. Shrubs to treelets, up to 2 m high; branchlets terete, c. 3 mm diam, densely hirsute, dark-brown when dry. Leaves: blades oblong-elliptic, rarely obovate, 10–15 by 4–6 cm, coriaceous, glabrous above except hirsute hairs on midrib, hirsute beneath, apex cuspidate to cuspidate-caudate, base cuneate, slightly asymmetric, nerves 9–13 pairs, flat to slightly depressed above, prominent beneath, ascending at an angle of c. 60°, curved to the margin, nervules obscure above, prominent beneath, subreticulate; petioles 5–8 mm long, densely dark-brown-hirsute. Stipules linear-lanceolate to triangular-lanceolate, up to 8 mm long, hirsute, cadulous. Cymes sessile; bracts numerous, hirsute, outer ones with a midrib, leaf-like, lanceolate, up to 1.5 cm long, hirsute, inner ones smaller, lanceolate to linear. Flowers sessile; calyx obconical, c. 2 mm long, densely longhirsute, lobes 5, subulate, c. 1 mm long; corolla tube c. 5 mm long, hirsute, lobes 5, triangular, c. 3 mm long, hirsute. Drupes subglobose, c. 3 mm diam, hirsute; pyrenes 5, smooth on the abaxial face. Distribution — Indonesia (Irian Jaya), Papua New Guinea. 37. Lasianthus cyanocarpus Jack Lasianthus cyanocarpus Jack (1823) 125; Miq. (1857) 316; (1869) 245 (p.p. excl. syn. L. densifolius Miq.); Merr. (1952) 229; Bakh.f. (1965) 338. — Type: Jack s.n. (lost), Sumatra, Tappanooly (west coast of Sumatra), Borssum 1864 (neo L, designated here), west coast, Lubuk Sikaping. Lasianthus inaequalis Blume (1826–1827) 996; Miq. (1857) 316; King & Gamble (1904) 114; Elmer (1906b) 10; Merr. (1921) 578; (1923) 566; Ridl. (1923a) 155; Valeton (1927) 106; Merr. (1937) 298; K.M.Wong (1989) 370; H.Zhu (2001b) 131, syn. nov. — Type: Blume s.n. (lecto L, here designated, barcode L0000688; isolecto K), Java, Seribu. Lasianthus cyanocarpus Jack var. subsessilis King & Gamble (1904) 113. — Syntypes: Ridley 9730 (n.v.), Perk; King’s collector 472 (K); Scortechini 1207 (K). Lasianthus subinaequalis King & Gamble (1904) 120; Ridl. (1923a) 155; Craib (1934) 218. — Syntypes: King’s collector 157 (n.v.), 3358 (n.v.), Ridley 9715 (SING), Wray 2588 (SING), 3451 (SING), Peninsular Malaysia, Perak. Lasianthus bractescens Ridl. (1918) 85; (1923a) 154; K.M.Wong (1989) 370, syn. nov. — Type: Ridley 8213 (syn K), Peninsular Malaysia, Selangor, Batu Caves. Lasianthus bractescens Ridl. var. rosulatus Ridl. (1918) 85; (1923a) 154, syn. nov. — Type: Ridley s.n. (n.v.), Peninsular Malaysia, Perak. Lasianthus inaequalis Blume var. angustibractea Valeton (1927) 107, syn nov. — Type: Schlechter 19334 (syn K), New Guinea. Shrubs; branches terete, 1.5–2 mm diam, densely tomentose. Leaves: blades elliptic-oblong to oblanceolate-oblong, 8–13.5 by 2.5–4.5 cm, membranaceous, glabrous except tomentose midrib and puberulous nerves above, or all tomentose, softly tomentose beneath, apex shortly acuminate or caudate, base rounded, oblique or oblique cordate, midrib conspicuous above, prominent beneath, nerves 7–10 pairs, flat above, prominent beneath, ascending at an angle of c. 50°, curved to the margin, nervules obscure above, distinct beneath, subreticulate; petioles short, c. 2 mm long, densely tomentose. Stipules linear, inconspicuous, densely tomentose, deciduous. Cymes sessile; bracts numerous, densely tomentose, outer ones leaf-like, broadly ovate, abruptly acuminate at apex, up to 1.8 cm long, long-tomentose, inner ones becoming smaller. Flowers sessile; calyx campanulate with 4–5 deep unequal lanceolate or linear-lanceolate tomentose 3–4 mm long lobes; corolla short, 4–5-lobed, glabrous inside, long yellow pilose outside. Drupes ovoid, sparsely pilose, crowned by the persistent calyx lobes; pyrenes 4, rugose. Distribution — Thailand, Malaysia (Peninsular, Borneo), Indonesia (Sumatra, Java, Kalimantan, Lesser Sunda Islands, Irian Jaya), Philippines, Papua New Guinea. 34 Blumea – Volume 57 / 1, 2012 Fig. 11 Lasianthus depressineurus H.Zhu. a. Habit with inflorescences; b. detail of lower surface of leaf; c. detail of upper surface of leaf showing reticulate nervules; d. detail of node showing stipule and axillary inflorescence; e. bract; f. bracteole; g. stipule; h. calyx split opened; i. immature fruit (all: Dyg Awa & B. Lee S.50579, holo L). 1 cm 2 mm 2 mm 3 mm 3 mm 3 mm 35 H. Zhu et al.: Revision of Malesian Lasianthus Notes — As mentioned earlier in this paper, Jack’s holos were destroyed by fire in 1824. The type of L. cyanocarpus was certainly lost. The name L. cyanocarpus has been misused for the species L. hirsutus for a long time and in many taxonomic references as pointed out by Merrill (1952). A specimen Blume s.n. (L0000688) from Seribu of Java indicated as L. inaequalis with Blume’s handwriting is recognised to be a syntype and here designated as lectotype because more than one locality was mentioned in Blume’s original description. Miquel (1869) reduced L. inaequalis to a synonym of L. cyanocarpus. Merrill (1952) confirmed that L. inaequalis from Java is conspecific to L. cyanocarpus. Bakhuizen van den Brink Jr (1965) followed Miquel and Merrill in the treatments of Java Lasianthus. From Jack’s description L. cyanocarpus has large cordate bracts, which is a key diagnostic character. After checking all Malesian specimens, we are sure that L. inaequalis is conspecific to L. cyanocarpus. Miquel’s and Merrill’s treatments are correct. Lasianthus cyanocarpus is very closely related to L. attenuatus, but differs from the latter by having cordate to broadly lanceolate, somewhere leaf-like bracts. Lasianthus cyanocarpus differs from L. hirsutus by having relatively small leaves with cordate, slightly oblique bases, usually less and relatively small bracts. Lasianthus hirsutus has relatively large leaves with cuneate, little unequal base; relatively long petioles; conspicuous triangular stipules; more bracts with outer ones ovate-oblong, and leaf-like, inner ones gradually narrower and smaller; as well as usually long rough hairs cover. From Ridley’s description, L. bractescens var. rosulatus has oblong, broad and sessile leaves with unequal base; bracts ovate acute, forming a rosette, which strongly suggest it is the same species as L. cyanocarpus although its type is not seen. 38. Lasianthus depressineurus H.Zhu, sp. nov. — Fig. 11 Proximus L. inodorus Blume, sed foliis minoribus ovato-lanceolatis, costis et nervis supra conspicue depressis, lobis calycis ovatis longioribus tubo differt. — Typus: Dyg Awa & Lee S.50579 (holo L; iso K, MO), Borneo, Sarawak, Bt. Lawi, Bario, 1290 m alt., in hill mixed dipterocarp forest, on top of the ridge, 3 Aug. 1985. Treelet, 2 m high; branchlets slender, terete, c. 1 mm diam, densely tomentose. Leaves: blades ovate-lanceolate, 8–11 by 1.5 cm, coriaceous, glabrous except pubescent midrib and nerves beneath, apex caudate, base acute or subrounded; midrib and nerves deeply depressed above, strongly prominent beneath, nerves 6–7 pairs, ascending at an angle of 45–60°, curved to margin, nervules obscure above, prominent beneath, reticulate, glabrous, margin conspicuously recurved; petioles 5–8 mm long, densely tomentose. Stipules triangular, c. 2 mm long, densely tomentose. Inflorescences sessile; bracts ovateorbicular, c. 3 mm long, pubescent surfaces; bracteoles oblong, c. 2.5 mm long, pubescent. Flowers fascicular, sessile; calyx campanulate, pubescent, lobes 5, ovate, c. 2.5 mm long, 1.5 mm broad, pubescent both surfaces, densely hirsute on margin, tubes c. 2 mm long; corolla not seen. Drupes globose, c. 3 mm diam, pubescent, crowned by persistent ovate, c. 4 mm long calyx lobes; pyrenes 4, smooth on the abaxial face. Distribution — Malaysian (Sarawak). Habitat & Ecology — In montane forest between 1290– 1400 m altitude. Flowering and fruiting in August. Additional specimen examined. Borneo, Sarawak, Limbang, D. Awa & B. Lee S.47839 (L). Note — This species is characterized by its deeply depressed midrib and nerves above surface and ovate-orbicular bracts. It seems to be related to L. inodorus, but conspicuously differs from the latter by having small ovate-lanceolate leaves with deeply depressed midrib and nerves above, ovate calyx lobes longer than the tube. 39. Lasianthus dinghouanus H.Zhu, sp. nov. — Fig. 12 Proximus L. laxifloroideo H.Zhu, sed foliis lineari-lanceolatis 1–2 cm latis apice longicaudatis, floribus minoribus, lobis calycis triangularis 0.8 mm longis differt. A L. foetidissimo A.Chev. ex Pit. foliis apice longicaudatis, nervis 3–4-binatis, floribus 4-meris, drupis glabris et pyrenis 4 differt. — Typus: Dyg Awa et al. S.46999 (holo L; iso K, L), Borneo, G. Kanji, Lindu, 530 m alt., in non-dipterocarp forest on sandy soil, 18 Apr. 1984. Treelet, 4 m high, 2 cm diam; branchlets slender, terete, c. 2 mm diam, subglabrous to appressed-pubescent, brown. Leaves: blades linear-lanceolate, 10–14 by 1–2 cm, chartaceous to subcoriaceous, glabrous except more or less appressed-pubescent midrib and nerves beneath, apex long caudate, margin recurved, base acute to acuminate, midrib flat above, prominent beneath, nerves flat above, prominent beneath, 3–4 pairs, nervules slightly obscure above, conspicuous beneath, parallel, glabrous; petioles 8–10 mm long, slender, more or less appressed-pubescent. Stipules subulate-triangular to triangular, 1–1.5 mm long, pubescent. Inflorescences peduncled; peduncles 5–20 mm long, slender, trifurcate at apex, more or less pubescent; bracts very small, subulate, c. 1 mm long, pubescent. Flowers shortly pediceled; calyx campanulate, pubescent, tube c. 1 mm long, lobes 4, triangular, c. 0.8 mm long; corolla c. 5 mm long, tube glabrous outside, lobes 4, ovate, c. 1 mm long, pubescent external surface. Drupes with 1–2 mm long pedicels, globose, c. 4 mm diam, glabrous but very verrucose, 4-ridged; pyrenes 4, verrucose on the abaxial face. Distribution — Endemic to Borneo. Habitat & Ecology — Lower montane forest between 500– 1300 m altitude. Flowering and fruiting between April and October. Additional specimens examined. Sarawak, Lundu, Ftr. Ilias Paie S.13619 (K, L); same locality, J.D. Mamit S.35169 (K, L); same locality, P.S.Ashton S.18876 (K, L); Sarawak, Gunong Silantek, Ilias Paie S.42455 (K, L); Sarawak, G.Buri, P.Chai et al. S.36803 (L). Note — This species is closely related to L. laxifloroideus. It differs from the latter by having glabrous or sparsely appressed-pubescent on branches and leaf midrib beneath, leaves linear-lanceolate, less than 2 cm broad; leaf apex long caudate; flowers relatively small, calyx lobes triangular, c. 0.8 mm long. It is also similar to L. foetidissimus, but differs from the latter by having leaves with a long caudate apex, nerves 3–4 pairs, flowers 4-merous, drupes glabrous with 4 pyrenes. Lasianthus foetidissimus has leaves with an acuminate and mucronate apex; nerves 6 pairs; flowers 5-merous; drupes pubescent with 5 pyrenes. 40. Lasianthus ellipticus Wight Lasianthus ellipticus Wight (1846) 507; Hook.f. (1880) 182; King & Gamble (1904) 118; Ridl. (1923a) 159; K.M.Wong (1989) 372; H.Zhu (2001b) 126. — Type: Kew Distrib. 2934 (Griffith s.n.) (holo K), Peninsular Malaysia. Lasianthus venulosus Ridl. (1912) 23, non Wight (1846). — Syntypes: Chan Chu Kang 6146 (n.v.), Ridley 12542 (iso K), Bukit Panjang, Singapore. Lasianthus sordidus Ridl. (1911) 60. — Type: Ridley 14488 (iso K), Peninsular Malaysia, Ulu Temengoh. Lasianthus scalaris Craib (1933) 26; (1934) 216. — Type: Kerr 18809 (holo K; iso BK), Thailand, Puket. Lasianthus velutinus Ridl. (1920) 144; (1923a) 159; K.M.Wong (1989) 372, syn. nov. — Type: Ridley s.n. (holo SING; iso K), Peninsular Malaysia, Kelantan. Shrubs, 1–2 m high, branchlets densely hirsute to tomentose. Leaves: blades elliptic or oblong-elliptic, 10–15 by 5–7 cm, rigid chartaceous to subcoriaceous, usually glabrous above, appressed-hirsute on midrib, nerves and nervules beneath, apex acute to shortly caudate or cuspidate-acuminate, base broadly cuneate to subrounded, midrib slightly depressed above, 36 Blumea – Volume 57 / 1, 2012 Fig. 12 Lasianthus dinghouanus H.Zhu. a. Habit with inflorescences; b. lower surface of leaf showing parallel nervules; c. detail of node showing stipule and axillary inflorescence; d. fruit; e. cross section of fruit (all: Dyg Awa et al. S.46999, holo and iso L). 2 cm 2 mm 2 mm 37 H. Zhu et al.: Revision of Malesian Lasianthus Fig. 13 Lasianthus eriocalyx H.Zhu. a. Habit with inflorescences; b. detail of branch; c. immature flower; d. calyx split opened (all: F.S.P. Ng s.n. KEP-FRI 22027, holo L). 6 mm 2 cm 2 mm 38 Blumea – Volume 57 / 1, 2012 prominent beneath, nerves and nervules conspicuous or slightly prominent above, prominent beneath, nerves 7–9 pairs, ascending at an angle of c. 50°, curved to the margin, nervules parallel, margin recurved; petioles 0.8–1.7 cm long, densely hirsute. Stipules broadly triangular, up to 7 mm long, 4.5 mm broad, hirsute. Cymes sessile; bracts conspicuous, up to 9 mm long, linear or lanceolate, densely long-hirsute. Flowers sessile; calyx long-hirsute, tube c. 1 mm long, lobes 5 narrowly triangular, c. 1 mm long, hirsute outside; corolla tube c. 4–5 mm long, glabrous, corolla lobes 5, c. 2 mm long, long-hirsute outside, floccose inside. Drupes subglobose c. 5 mm diam, hirsute; pyrenes 5. Distribution — Thailand, Malaysia (Peninsular, Borneo), Indonesia (Sumatra, Java). Note — Lasianthus velutinus is almost the same as L. ellipticus except a little denser tomentose hairs which is not enough to separate it from the latter. 41. Lasianthus eriocalyx H.Zhu, sp. nov. — Fig. 13 Similis L. ferrugineus King & Gamble, sed foliis longissimis, stipulis minutis 1 mm longis et lobis calycis 7–8 differt. — Typus: F.S.P. Ng s.n. KEP-FRI 22027 (holo L; iso K), Peninsular Malaysia, Ulu Trengganu, 900 m alt., in hillside primary forest, 26 Mar. 1974. Treelet, c. 6 m high, crown spreading horizontally; branchlets terete, densely yellowish tomentose. Leaves: blades narrowly oblong-lanceolate, 23–25 by 3–4 cm, chartaceous, tomentose on both surfaces, apex acuminate to caudate, base acute to subrounded, midrib densely tomentose, prominent beneath, nerves flat above, prominent beneath, tomentose, 16–18 pairs, ascending at an angle of 60°, curved to margin, nervules obscure above, conspicuous beneath, reticulate, tomentose; petioles c. 5 mm long, tomentose. Stipules small, triangular, c. 1 mm long, tomentose. Inflorescences sessile; bracts absent. Flowers in fascicle, sessile; calyx densely tomentose, tube c. 2.5 mm long, lobes 7–8, lanceolate, 3 mm long, 1 mm broad, densely tomentose external surface, sericeous internal surface; corolla not seen (from the description on the label of the type specimen, the corolla is whitish green, filled with hairs inside). Fruits not seen. Distribution — Peninsular Malaysia. Habitat & Ecology — In montane primary forest. Flowering in March. Note — This species is characterized by having very tomentose hairs on branchlets, leaves and calyx; leaves narrowly oblong-lanceolate, very long; calyx with 7–8 lanceolate lobes. This species is most similar to L. ferrugineus, but differs from the latter by having much longer leaves (23–25 cm long), very small stipules (1 mm long) and calyx lobes 7–8. Lasianthus ferrugineus has leaves less than 18 cm long, stipules more than 3 mm long and calyx lobes 5. 42. Lasianthus euneurus Stapf Lasianthus euneurus Stapf (1894) 185; Merr. (1921) 577. — Type: Haviland 1119 (holo K; iso SING), Borneo, 7600 ft, in upper montane rainforests. Shrubs to treelets, 2 m high; branchlets terete, c. 2 mm diam, densely pubescent. Leaves: blades elliptic or oblong-elliptic, 5–10 by 1.5–4 cm, coriaceous, glabrous except pubescence on midrib and nerves beneath, apex acute to acute-cuspidate, base acute, midrib and nerves flat or slightly prominent above, strongly prominent beneath, nerves 7–8 pairs, ascending at an angle of 50–60°, curved to the margin, nervules distinct and nitid both surfaces, densely anastomosing, parallel; petioles 5–10 mm long, densely pubescent. Stipules subulate-triangular, c. 1 mm long, pubescent. Cymes sessile; bracts absent. Flowers sessile; calyx obconical to campanulate, c. 1.5 mm long, pubescent outside, calyx limb truncate or minutely 4-toothed; corolla tube c. 3–4 mm long, puberulous outside, corolla lobes 4, ovate, pubescent outside, floccose inside; anthers 4. Drupes subglobose, 2–3 mm diam, sparsely pubescent, 4-obtuseridged; pyrenes 4. Distribution — Malaysia (Borneo) and Indonesia (Sumatra). 43. Lasianthus fasciculus H.Zhu, sp. nov. — Fig. 14 Proximus L. euneuro Stapf, sed floribus 5-meris, calycibus 3–4 mm longis, lobis calycis triangulatis 1–1.5 mm longis, foliis 10–15 cm longis, nervis 9–11-binatis et nervulis supra obscuris differt. — Typus: Kessler PK32 (holo L), Sumatra, Kabulaten Tapanuli Utara Tele, 1650 m alt., in primary forest, 2 Dec. 1988. Treelet, c. 4–5 m high; branchlets compressed at the youngest part, densely appressed-pubescent. Leaves: blades oblong, 10–15 by 3–5 cm, subcoriaceous, glabrous above, pubescent venation beneath, apex acute, mucronate, base acute, margin recurved, midrib depressed above, strongly prominent beneath, nerves flat or slightly depressed above, prominent beneath, 9–11 pairs, ascending at an angle of 60–70°, curved to margin, nervules obscure above, prominent beneath, parallel; petioles 1–1.5 cm long, densely appressed-pubescent. Stipules triangular, c. 1 mm long, densely appressed-pubescent. Inflorescences sessile; bracts inconspicuous. Flowers fascicular, sessile; calyx campanulate, 2–4 mm long, densely pubescent, lobes 5, triangular, 1–1.5 mm long, pubescent except margins external surface, tube 1.5–2 mm long, densely pubescent; corolla 6–7 mm long, lobes 5, triangular-lanceolate, c. 2 mm long, densely pubescent external surface, floccose internal surface, tube glabrous at the lower part. Fruits not seen. Distribution — Indonesia (West Sumatra). Habitat & Ecology — In montane forests between 1650– 2300 m altitude. Flowering during December and February. Additional specimen examined. W Sumatra, Gunung Rasam, H. Nagamasu 3691 (L). Note — This species is closely related to L. euneurus, but differs from the latter by having 5-merous flowers, calyx 3–4 mm long with triangular, 1–1.5 mm long lobes, leaves relatively big, 10–15 by 3–5 cm, with 9–11 pairs of nerves, nervules obscure above. Lasianthus euneurus has 4-merous flowers, calyx 1–1.5 mm long with 4 minute teeth, leaves less than 10 cm long with 7–8 pairs of nerves, nervules conspicuous and prominent above. 44. Lasianthus ferrugineus King & Gamble Lasianthus ferrugineus King & Gamble (1904) 130. — Type: Wray 3911 (holo SING; iso K), Peninsular Malaysia, Perak, 4500 ft. Shrubs to treelets; c. 2 m high; branchlets terete, robust, c. 3 mm diam, densely brown-tomentose. Leaves: blades oblonglanceolate, 12–15 by 2.5–3.5 cm, subcoriaceous, tomentose on both surfaces but denser beneath, apex acuminate to caudate-acuminate, base broadly cuneate to subrounded, slightly asymmetric, dark-brown when dry, midrib dense browntomentose on both surfaces, prominent beneath, nerves 14–18 pairs, flat above, prominent beneath, ascending at an angle of c. 70°, curved to the margin, nervules obscure above, distinct beneath, reticulate; petioles c. 5 mm long, densely browntomentose. Stipules triangular, 3–4 mm long, densely browntomentose. Cymes sessile, bracts absent. Flowers sessile; calyx tubular-campanulate, densely tomentose, c. 3 mm long, with 5 lanceolate or subulate lobes; corolla not seen. Drupes subglobose, densely brown-tomentose, 3–4 mm diam; pyrenes 5–6, verrucose on the abaxial face, thick-walled. Distribution — Peninsular Malaysia. 39 H. Zhu et al.: Revision of Malesian Lasianthus Fig. 14 Lasianthus fasciculus H.Zhu. a. Habit with inflorescences; b. lower surface of leaf showing parallel nervules; c. detail of node showing stipule and axillary inflorescences; d. calyx split opened; e. corolla split opened (all: P.J.A. Kessler PK32, holo L). 2 cm 2 mm 2 mm 40 Blumea – Volume 57 / 1, 2012 Fig. 15 Lasianthus filipedunculatus H.Zhu. a. Habit with inflorescences; b. detail of lower surface of leaf showing subparallel nervules and indumentum; c. detail of node; d. flower; e. fruit; f. cross section of fruit (all: I.M. Said et al. BRUN15640, holo L). 4 mm 2 cm 2 mm 8 mm 41 H. Zhu et al.: Revision of Malesian Lasianthus Habitat & Ecology — In upper montane rainforest. Flowering and fruiting in March and April. Notes — This species is similar to L. pilosus, especially in the indumentum. Wong (1989) reduced L. ferrugineus to a synonym of L. pilosus. However, it differs from the latter by having leaves with broad-cuneate bases, longer petioles and shorter calyx lobes. We think it is better to keep it as a separate species. This species differs from L. hirtus by having relatively small triangular stipules, oblong-lanceolate leaves 12.5–15 cm long, with 14–18 paired-nerves. From King, it differs from L. harveyanus King & Gamble by having leaves sparsely hairy above, shorter hairy beneath, oblong-lanceolate, nerves 16–18; calyx with 5 deep lanceolate lobes; while the latter has leaves scabrid puberulous, elliptic to ovate-elliptic, nerves 4–6 pairs, calyx narrowly campanulate. 45. Lasianthus filiformis King & Gamble Lasianthus filiformis King & Gamble (1904) 131; Ridl. (1923a) 169; K.M.Wong (1989) 367. — Type: Scortechini 29 (lecto SING, designated by Wong 1989), Peninsular Malaysia, Perak; King 857 (para K), Peninsular Malaysia, Perak. Lasianthus filiformis King & Gamble var. bracteata King & Gamble (1904) 131; K.M.Wong (1989) 367. — Type: Ridley 8578 (lecto SING, designated by Wong 1989), Peninsular Malaysia, Selangor; Scortechini 190 (para K), Peninsular Malaysia, Perak. Slender shrubs, c. 1 m high; branchlets slender, c. 1.5 mm diam, spreading long-hairy. Leaves: blades oblong to oblonglanceolate, 5–7 by 1.5–2 cm, chartaceous, villous on both surfaces, apex acuminate or cuspidate-acuminate, base rounded to subcordate, oblique, midrib and nerves slender and slightly depressed above, prominent beneath, nerves 6–9 pairs, ascending at an angle of c. 45°, curved to the margin; nervules obscure above, distinct beneath, subparallel; petioles very short, c. 1 mm long, villous. Stipules linear, c. 4 mm long, villous. Cymes pedunculate; peduncles slender, filiform, 1–2.5 cm long, villous; bracts 1–2, leaf-like and ovate, or linear-lanceolate, 6–10 mm long, villous. Flowers sessile; calyx campanulate, villous, tube c. 2 mm long, lobes 4, linear-lanceolate, c. 2 mm long; corolla tubular, villous, 4-lobed; ovary 8-locular. Drupes globose, c. 3 mm diam, sparsely long-hairy; pyrenes 6–8. Distribution — Peninsular Malaysia. Note — This var. bracteata differs from the typical variety by having only a little wider bracts which is not enough to recognize it as a variety. 46. Lasianthus filipedunculatus H.Zhu, sp. nov. — Fig. 15 A L. pendulo Ridl. ramulis foliis subtus pedunculis bracteis molliter tomentosis, costis et nervis supra leviter depressis vel planiusculis, petiolis 5 mm longis, capitulis cymae laxis, floribus 4-meris et pyrenis 4 differt. — Typus: BRUN 15640 (holo L), Brunei, Temburong, in primary dipterocarp forest, 17 July 1993. Treelet; branchlets terete, robust, c. 2–3 mm diam, brownyellowish tomentose. Leaves: blades oblong-ovate to ovate, 10–15 by 2.5–5 cm, chartaceous to subcoriaceous, glabrous above, tomentose beneath, apex caudate to cuspidate, margin recurved, base rounded to subcordate, midrib flat to slightly depressed above, prominent beneath, 4–6 pairs, ascending at an angle of c. 60°, curved to margin, nervules obscure above, conspicuous beneath, subreticulate, tomentose beneath; petioles c. 5 mm long, densely tomentose. Stipules small, triangular, less than 2 mm long, densely tomentose. Inflorescences pedunculate; peduncles slender, 4–13 cm long, tomentose; bracts 2 in each node, thinly linear, 5–20 mm long, tomentose; bracteoles linear, relatively short, tomentose. Flowers 1–2 in each node, 4-merous, pediceled; pedicels c. 1 mm long; calyx densely tomentose, lobes 4, linear, c. 2.5 mm long, tube short, obconical, c. 1 mm long; corolla c. 3 mm long, upper part external surface tomentose, lower part glabrous; stamens 4. Drupes subglobose, 7–8 mm diam, square in transverse section, warty, sparsely tomentose, conspicuously 4-ridged, crowned by 4 linear tomentose 5 mm long calyx lobes; pyrenes 4, deeply sulcate on the abaxial face, subrhombic in transverse section. Distribution — Endemic to Borneo: Brunei, Sabah. Habitat & Ecology — In primary forest of lower montane below 1200 m altitude. Flowering in July and fruiting in November. Additional specimens examined. Sabah, Nabawan, F. Krispinus SAN 118556 (K, L), F. Krispinus SAN 118646 (K, L); Sabah, Pensiangan, A. Mantor SAN 136779 (L); Brunei, Temburong, D.A. Simpson 2201 (K, L), same locality, P.C. Bygrave 35 (L). Notes — This species differs from L. pendulus by having soft and relatively short hairs on branches, leaves beneath, peduncles and bracts; leaf midrib and nerves slightly depressed or flat above, petioles c. 5 mm long; head of cymes less compact; flowers 4-merous; drupes with 4 pyrenes. Lasianthus pendulus has relatively long and stiff hairs on branches, leaves beneath, peduncles and bracts; leaf midrib and nerves deeply depressed above; petioles 1–2 mm long, head of cymes more compact; flowers 5-merous; drupes with 5 pyrenes. This species is also related to L. robinsonii, but differs in having oblong-ovate to ovate leaves, nerves 4–6 pairs, petioles c. 5 mm long and stipules triangular less than 2 mm long. Lasianthus robinsonii has oblong leaves, nerves 8–9 pairs, petioles 2–3 mm and stipules triangular lanceolate, 4–5 mm long. 47. Lasianthus flavinervius Ridl. Lasianthus flavinervius Ridl. (1915) 154; (1923a) 162. — Type: Ridley 16121 (holo SING; iso K), Peninsular Malaysia, Gunong Tahan, Pahang. Shrubs; branchlets robust, terete, 3–4 mm diam, compressed and slightly sulcate in the youngest part, densely appressedpubescent. Leaves: blades oblong to oblong-elliptic, 10–15 by 3–5 cm, coriaceous, glabrous above, appressed-pubescent on venation beneath, apex acuminate to cuspidate-acuminate, base acute or cuneate; midrib depressed above, strongly prominent beneath, nerves 6–8 pairs, flat above, strongly prominent beneath, ascending at an angle of 50–60°, curved to the margin, nervules slightly prominent above, conspicuously prominent beneath, subparallel; petioles c. 1 cm long, densely appressed-pubescent. Stipules broadly triangular, c. 1.5 mm long, appressed-pubescent. Cymes sessile; bracts absent. Flowers sessile; calyx tubular-campanulate, 3–4 mm long, appressed-pubescent outside, lobes 5, ovate-triangular, curved inward, c. 1–1.5 mm long, hirsute inside; corolla hirsute outside. Drupes campanulate, c. 5 mm long, crowned by 5 calyx lobes, pubescent, conspicuously 4–5-ridged; pyrenes 4–5. Distribution — Peninsular Malaysia, Indonesia (Sumatra). Notes — The species is very close to L. chinensis. It differs from the latter by having 6–8 pairs of nerves and somewhere subparallel nervules, midrib depressed above and strongly prominent beneath, nerves strongly prominent beneath. Wong (1989) sank this species to L. perakensis. However, it differs from the latter by having midrib and nerves strongly prominent beneath, calyx tubular-campanulate with 5 ovatetriangular lobes, drupes campanulate crowned by 5 conspicuous calyx lobes, drupes with 4–5 ridges. Lasianthus perakensis has midrib and nerves not strongly prominent beneath, calyx limb slightly trumpet-shaped with 5 indistinct obtuse lobes, drupes hemispheric with 10 obtuse ridges and crowned by obtuse calyx lobes. 48 Blumea – Volume 57 / 1, 2012 nerves flat above, prominent beneath, 8–11 pairs, ascending at an angle of 70–80°, curved to the margin, nervules obscure above, slightly clear beneath, reticulate, puberulous beneath; petioles 1.2–1.5 cm long, glabrous. Stipules triangular, c. 2 mm long, hirsute. Inflorescences sessile; bracts inconspicuous. Flowers sessile; calyx pubescent, with 4–5 minutely teeth; corolla pubescent. Drupes c. 5 mm diam, warty, irregular ridged, pubescent; pyrenes 4–5. Distribution — Indonesia (Sumatra). Habitat & Ecology — In lower montane forests between 1100–1500 m altitude in W Sumatra. Flowering and fruiting between January and August. Additional specimen examined. W Sumatra, Padang, H. Nagamasu 3140 (L). Note — This species is similar to L. angustifolius. It differs from the latter by having glabrous branchlets, densely hirsute leaf margin, nerves 8–11 pairs and longer petioles (1.2–1.5 cm long); while L. angustifolius has pubescent branchlets, glabrous leaf margin, nerves 12–14 pairs and short petioles (4–5 mm long). 58. Lasianthus hirtus Ridl. Lasianthus hirtus Ridl. (1909) 36; (1923a) 153. — Type: Ridley 14009 (holo SING; iso K), Peninsular Malaysia, Pahan. Lasianthus acutatus auct non Miq.: Bakh.f. (1965) 341. Shrubs, 2 m high; branchlets terete, c. 2 mm diam, densely spreading brown-pilose. Leaves: blades oblong-lanceolate, 6–9 by 2–2.5 cm, thinly coriaceous, densely pilose on both surfaces, apex caudate to caudate-acuminate, base acute to subrounded, midrib and nerves depressed above, prominent beneath, nerves 8–9 pairs, ascending at an angle of c. 50°, curved to the margin, nervules obscure above, distinct beneath, subparallel; petioles 2–4 mm long, densely spreading brown-pilose. Stipules ovate, 3–4 mm long, membranaceous, densely pilose. Cymes sessile; bracts absent. Flowers sessile; calyx campanulate, dense pilose, tube c. 1 mm long, lobes 5, subulate-lanceolate, c. 1.5 mm long, densely long-pilose; corolla c. 7–8 mm long, pilose outside, lobes 5, lanceolate. Drupes ellipsoid, c. 4 mm long, sparsely pilose to densely pilose, with 5 obtuse ridges, crowned by densely pilose calyx limbs; pyrenes 4, smooth on the abaxial face. Distribution — Peninsular Malaysia, Indonesia (Sumatra, Java, Sulawesi). Notes — This species is close to L. acutatus but differs from it by having dense spreading pilose hairs on branches, leaves, petioles, stipules and calyx; leaves relatively small (6–9 cm long); stipules ovate. Lasianthus acutatus has stiff hairs on branches, leaves, and stipules, relatively bigger leaves that are 12–17 cm long, linear-lanceolate stipules. Lasianthus hirtus has ovate and membranaceous stipules, the same as L. stercorarius, but differs from the latter by having dense spreading pilose hairs on branches, leaves, petioles, stipules, and calyx; calyx with 5 subulate-lanceolate lobes; drupes with 5 pyrenes. Lasianthus hirtus was reduced to a synonym of L. pilosus by Wong (1989). However, it conspicuously differs from L. pilosus by having leaf blades relatively small (6–9 cm long) and with less nerves and acute to subrounded base; stipules ovate; bracts absent; calyx lobes subulate-lanceolate, relatively short, c. 1.5 mm long. Lasianthus pilosus has leaves relatively large (10–17 cm long) with usually rounded unequal base and more nerves; stipules triangular-lanceolate; bracts subulate; calyx lobes filiform, relatively long. Lasianthus hirtus could be conspecific to L. rufus (Korth.) Miq., because it almost agrees with Korthals’ (1851) description. Without seeing type specimens of L. rufus, we still treat L. hirtus separately. 59. Lasianthus hispidulus (Drake) Pit. — Map 6 Lasianthus hispidulus (Drake) Pit. (1924) 393; H.Zhu (2001b) 130; (2002) 99. — Mephitidia hispidulus Drake (1895) 239. — Lectotype (selected by Zhu 2001b): Eberhardt 3852 (lecto P), Vietnam. Thai-nguyen. Lasianthus brevidens Craib (1933) 19; (1934) 208; H.Zhu (1998) 151. — Type: Kerr 10833 (holo K; iso BM), Thailand, Rachaburi, Prachuap, 700 m. Lasianthus bunzanensis Shimizu (1944) 301; T.S.Liu & J.M.Chao (1964) 121, f. 3; H.Zhu (1994) 73; H.J.Li & H.Y.Liu (1998) 277, pl. 122; H.S.Lo (1999) 99. — Type: Suzuki 16385 (holo TIU, n.v.), Taiwan, Taipei. Shrubs, 1–1.5 m high; branchlets terete, c. 2 mm diam, densely brown-villous. Leaves: blades elliptic to ovate-oblong, 7–10 by 3–5 cm, thinly coriaceous to chartaceous, apex cuspidateacuminate or acuminate, base acute or obtuse, glabrous above, densely villous beneath, midrib and nerves flat above, prominent beneath, nerves 4–6 pairs, ascending at an angle of 60–70°, curved to the margin, nervules slightly obscure above, distinct beneath, subparallel; petioles 4–7 mm long, villous. Stipules minute, subulate, c. 1 mm long, hirsute. Cymes sessile; bracts absent. Flowers sessile, clusters; calyx campanulate, 1.5 mm long, densely villous, teeth 5, minute, broad-triangular, c. 0.5 mm long, densely villous outside, glabrous inside; corolla white, 5–6 mm long, 5-lobed, pilose outside, villous inside. Drupes globose, hirsute, 3–5 mm diam; pyrenes 5. Distribution — China, Japan (Ryukyu), Vietnam, Thailand, Malaysia (Borneo), Indonesia (Sumatra, Java, Irian Jaya) and Papua New Guinea. Note — Lasianthus brevidens from Thailand agrees well with L. hispidulus from Indochina, China (Hainan and Yunnan). We have not seen the type of L. bunzanensis from Taiwan, but from the original description and the checked specimens from Taiwan it is clear that L. bunzanensis is conspecific to L. hispidulus. This species is very closely related to L. clementis from the Philippines. It is separated from L. clementis only by having densely spreading hairs on branchlets, leaf undersurface, calyx and drupes, while L. clementis has densely more or less appressed hairs on branchlets, leaves beneath, and calyx, drupes appressed-pubescent or subglabrous. Both L. hispidulus and L. clementis have close affinity to L. fordii. They differ from L. fordii only by being densely hairy on branches, leaves beneath, and calyx. 60. Lasianthus inodorus Blume Key to the subspecies 1. Branchlets and leaves glabrous or sparsely puberulous hairs on leaf lower surface; leaves relatively large, longer than 12 cm ..........................................2 1. Branchlets and leaf lower surfaces appressed-pubescent; leaves relatively small, usually less than 12 cm long .... ............................. c. subsp. pubescens 2. Branchlets with conspicuous lenticel; leaves oblong with subrounded bases; nerves 7–11 pairs; nervules subreticulate; bracts caducous............b. subsp. montigenus 2. Branchlets without conspicuous lenticel; leaves elliptic to elliptic-lanceolate with acuminate bases; nerves 6–7 pairs; nervules parallel; bracts persistent ... a. subsp. inodorus a. subsp. inodorus — Map 8 Lasianthus inodorus Blume (1826–1827) 998; Miq. (1857) 322; (1869) 248; Merr. (1921) 577; Bakh.f. (1965) 339; H.Zhu (1998) 152; (2001a) 451; (2001b) 132; (2002) 77. — Mephitidia inodora (Blume) DC. (1830) 453. — Type: Blume s.n (holo L, barcode L0000690), Java. 49 H. Zhu et al.: Revision of Malesian Lasianthus Lasianthus tubiferus Hook.f. (1880) 183; Craib (1934) 219; Deb & M.Gangop. (1991) 305, f. 14; H.Zhu (1994) 59. — Type: Hooker & Thomson s.n. (lecto K; designated by Deb & Gangopadhyay 1991; isolecto A, L), India, Khasia. Lasianthus poilanei Pit. (1924) 398. — Type: Poilane 234 (holo P; iso E, K), Mt Elephant, 1000 m, Cambodia. Shrubs, 2–3 m high; branches and branchlets glabrous or thinly puberulous. Leaves: blades elliptic to elliptic-lanceolate, 10–20 by 2.5–6 cm, coriaceous, glabrous above, glabrous or sparsely puberulous beneath, apex acuminate, base acuminate; nerves 6–7 pairs, nervules parallel, nerves and nervules conspicuous beneath; petioles 6–10 mm long, glabrous or sparsely puberulous. Stipules triangular or ovate-lanceolate, 3–5 mm long, glabrous, coriaceous. Cymes sessile; bracts persistent, 3–6 mm long, orbicular or ovate-orbicular, thickly coriaceous, glabrous or pubescent. Flowers sessile; calyx tubular, tube 2 mm long, glabrous, with 5 ovate-lanceolate lobes, puberulous outside; corolla 10 mm long, puberulous outside, villous inside. Drupes ovoid, 10 mm long, glabrous; pyrenes 5. Distribution — India (northeast part), Bangladesh (eastern part), China (Yunnan), Vietnam, Cambodia, Thailand, Indonesia (Sumatra, Java). Note — Subsp. inodorus has a wide, disjunct distribution. However, the specimens from mainland SE Asia and the ones from Sumatra and Java are almost identical so that clear variation within this subspecies cannot be recognized. b. subsp. montigenus H.Zhu — Map 8 Lasianthus inodorus Blume subsp. montigenus H.Zhu (2001a) 451. — Type: Zhu 98-7-01 (holo HITBC), Borneo, Kinabalu, alt. 2400 m, in upper montane forest. Shrubs, 2–3 m tall; branches subterete or slightly tetragonous, striate and conspicuously lenticellate, glabrous, incrassate at nodes, internode 3.5–4 cm long, nigrescent when drying. Leaves: blades 12–15 by 3.5–5.5 cm, oblong, apex shortly caudate, base subrounded, coriaceous, glabrous on both sides, nerves 7–11-paired, tertiary nerves subreticulate, nerves and tertiary nerves conspicuous on both sides; petioles 5 mm long, glabrous. Stipules ovate-triangular, 3–5 mm long, apex acuminate, pubescent. Inflorescences sessile; bracts orbicular, subcoriaceous, caducous. Flowers sessile; calyx glabrescent, tube 2.5 mm long, lobes 5, broad-triangular, 2 mm long, apex and margin pubescent. Fruits oblong, 10 mm long, glabrous; pyrenes 5. Distribution — Endemic to Borneo. Additional specimens examined. Borneo, Kinabalu, alt. 7000 ft, Clemens 32536 (K). c. subsp. pubescens H.Zhu — Map 8 Lasianthus inodorus Blume subsp. pubescens H.Zhu (2001a) 451. — Type: Zhu 98-7-02 (holo HITBC), Borneo, Kinabalu, alt. 1600 m, in lower montane forest. Shrubs, 1–1.5 m tall; branches subterete, densely appressedpubescent, incrassate at nodes, internode 2.5–3 cm long. Leaves: blades 6–13 by 2.2–5 cm, oblong-lanceolate, apex subacute to shortly acuminate, base broadly cuneate, coriaceous, glabrous above, pubescent beneath, nerves 6–7-paired, tertiary nerves subparallel, nerves and tertiary nerves conspicuous on both sides; petioles 3 mm long, pubescent. Stipules triangular, 5 mm long, apex acuminate, densely pubescent. Inflorescences sessile; bracts orbicular, subcoriaceous, pubescent, semi-persistent. Flowers sessile; calyx tube 2 mm long, pubescent outside, lobes 5, 2 mm long, broad-triangular, pubescent outside, apex and margin densely pubescent; corolla 8 mm long, tube 4–5 mm long, glabrescent outside, lobes subspathulate, pubescent outside, villous inside; stamens 5; anthers oblong. Mature fruit not seen. Distribution — Endemic to Borneo (Sabah, Brunei, Sarawak, Kalimantan). Notes — Subsp. pubescens has coriaceous elliptic or ellipticlanceolate leaves and conspicuously orbicular or ovate coriaceous bracts that make it a clear species in the genus. Lasianthus inodorus from the mountains of Sumatra and Java is clearly conspecific to L. poilanei of Cambodia and L. tubiferus from India, Thailand and Yunnan of China. The species has coriaceous elliptic or elliptic-lanceolate leaves and persistent orbicular coriaceous bracts, which clearly distinguish it in the genus. 61. Lasianthus iteoides Valeton ex H.Zhu, sp. nov. — Fig. 18 Similis L. hexandro Blume, sed lobis calycis triangulatis c. 1 mm longis aequalis tubo differt. — Typus: Bünnemeijer 3578 (holo L), Indonesia, Sumatra Barat, Malintang, Gunung, c. 1100 m alt., 18 July 1918. Shrub, c. 3 m high; branchlets terete, c. 1.5 mm diam, densely appressed hirsute. Leaves: blades lanceolate to oblong lanceolate, 6–13 by 1.6–3 cm, subcoriaceous, glabrous above except hirsute midrib, villose beneath; apex acute to acuminate, margin slightly recurved, base acute, midrib depressed above, Map 8 Distribution of Lasianthus inodorus Blume (▲), L. inodorus subsp. montigenus H.Zhu (●) and L. inodorus subsp. pubescens H.Zhu (¡). 50 Blumea – Volume 57 / 1, 2012 Fig. 18 Lasianthus iteoides Valeton ex H.Zhu. a. Habit; b. detail of lower surface of leaf showing indumentum; c. detail of node showing axillary inflorescences; d. calyx split opened; e. corolla; f. fruit; g. cross section of pyrene (all: Bünnemeijer 3578, holo L). 1 mm 2 cm 2 mm 2 mm 5 mm 51 H. Zhu et al.: Revision of Malesian Lasianthus prominent beneath, hirsute, nerves depressed and obscure above, prominent and conspicuous beneath, hirsute beneath, 7–8 pairs, ascending at an angle of 60–70°, curved to margin, nervules obscure above, conspicuous beneath, villose, reticulate; petioles c. 5 mm long, densely appressed hirsute. Stipules triangular, 1 mm long, appressed hirsute. Inflorescences sessile; bracts inconspicuous. Flowers fascicular, sessile; calyx obconical, c. 2 mm long, densely villose, lobes 4–5, triangular, 1 mm long, villose both surfaces; corolla claviform when in bud, densely spreading villose, 6–7 mm long, lobes triangular-oblong, c. 2 mm long; stamens with c. 0.8 mm long filaments; styles 5 mm long. Drupes globose, c. 5 mm diam, villose; pyrenes 5, smooth on the abaxial face. Distribution — W Sumatra. Habitat & Ecology — In forest between 100–1100 m altitude. Flowering in July; fruiting in September. Additional specimens examined. Sumatra, Malintang, Bünnemeijer 3862 (L); Sumatra, Sago, Gunung, Bünnemeijer 3686 (L), Dayar Arbain DA399 (L). Notes — This species is similar to L. hexandrus, but differs from the latter by having triangular calyx lobes, c. 1 mm long, as long as the calyx tube. Lasianthus hexandrus has lanceolate calyx lobes, 3–4 mm long, conspicuously longer than the tube. Valeton recognized the species as a new one, but did not made the description except a determination label on the type specimen. Here we described it as a new species. 62. Lasianthus iteophyllus Miq. Lasianthus iteophyllus Miq. (1869) 247; Bakh.f. (1965) 340. — Lectotype (designated here): Blume s.n. (lecto L, barcode L0305214). Paratypes: Blume s.n. (L, barcode L0305211), Blume s.n. (L, barcode L0305213), Java. Shrubs; branchlets terete, c. 2 mm diam, sparsely hirsute, darkbrown in the youngest part. Leaves: blades lanceolate, 12–14 by 2.5–3 cm, thinly coriaceous, glabrous on both surfaces except sparsely hirsute hairs on midrib and nerves beneath, apex caudate-acuminate, base acute, slightly asymmetric, midrib and nerves slightly prominent above, prominent beneath, nerves 10–11 pairs, ascending at an angle of 60–70°, curved to the margin, nervules distinct and prominent on both surfaces, reticulate; petioles 3–4 mm long, hirsute. Stipules triangular, 3–4 mm long, glabrous except long-pilose hairs on the margin, coriaceous. Cymes sessile; bracts absent. Flowers sessile; calyx campanulate, 2–2.5 mm long, glabrous, limb pilose inside, with 5–6 minute teeth, teeth subulate, c. 0.5 mm long, pilose at the margin and inside; corolla not seen. Drupes globose, glabrous, very verrucose; pyrenes 5–6, irregular sulcate on the abaxial face. Distribution — Indonesia (Java). 63. Lasianthus kinabaluensis Stapf Lasianthus kinabaluensis Stapf (1894) 184; Wernham ex Gibbs (1914) 95; Merr. (1921) 577. — Type: Haviland 1306 (lecto K; designated here; isolecto SING); Haviland 1323 (para K), Borneo, Penokok, 3000–3200 ft. Shrubs, up to 2 m high, glabrous; branchlets terete, slender, c. 1 mm diam, nitid. Leaves: blades lanceolate, 5–10 by 1.5–2.5 cm, thinly coriaceous, apex caudate-acuminate, base acute or cuneate, the margin often reflexed, midrib flat or depressed above, prominent beneath, nerves slender, prominent beneath, 3–4 pairs, ascending at an angle of over 60°, curved near the leaf margin and joining the veins above forming an intramarginal vein, nervules slender, loosely anastomosing, distinct beneath, reticulate; petioles 5–10 mm long. Stipules minute, triangular, c. 1 mm long, ciliate at margin. Cymes sessile; bracts absent. Flowers sessile; calyx campanulate, c. 1.5 mm long, glabrous, lobes 4–5, triangular, c. 0.8 mm long; corolla c. 4–5 mm long, glabrous outside, lobes 4–5. Drupes subglobose, c. 5 mm diam, glabrous, 3–4-obtuse-ridged, verrucose; pyrenes 3–4, verrucose on the abaxial face. Distribution — Endemic to Borneo. Note — This species is related to L. constrictus by its glabrous plants, calyx constricted at the base of the limb and warty drupes with 3–4 pyrenes. It differs from the latter by having relatively small and thin leaves with looped venation. 64. Lasianthus laevigatus Blume Lasianthus laevigatus Blume (1826–1827) 998; Miq. (1857) 320; (1869) 247; Boerl. (1891) 140; Merr. (1921) 577; Bakh.f. (1965) 337. — Lectotype (designated here): Blume s.n. (lecto L, barcode L0000692). Paratypes: Blume s.n. (L, barcode L0000691), Blume s.n. (L, barcode L0000693), Blume s.n. (L, barcode L0000694), Java, Salak. Shrubs, up to 5 m high, glabrous; branchlets terete, 2–3 mm diam, dark-brown when dry. Leaves: blades oblong-lanceolate, 9–18 by 2–4.5 cm, coriaceous, apex cuspidate-acuminate, margin often reflexed and more or less undulate, base acuminate, midrib and nerves prominent on both surfaces, nerves 8–10 pairs, slender, ascending at an angle of over 60°, curved to the margin, nervules obscure above, distinct beneath, reticulate; petioles 1–2 cm long. Stipules triangular, c. 3 mm long, glabrous. Cymes pedunculate, glabrous; main peduncles 1–1.5 cm long; branches 5–8 mm long; bracts subulate, 1–2 mm long, minutely ciliate. Flowers sessile; calyx tube tubularcampanulate, c. 5 mm long, glabrous, calyx lobes 4, lanceolate, 5–6 mm long, glabrous; corolla 7–8 mm long, tube glabrous, lobes 6, glabrous outside, densely floccose inside; stamens 6. Drupes turbinate, c. 5–6 mm diam, glabrous, conspicuously 6-ridged; pyrenes 6. Distribution — Endemic to Indonesia (Java). Habitat & Ecology — From lowland rainforests to montane forests between 700–1400 m altitude. Flowering in March; fruiting in April. 65. Lasianthus lancilobus H.Zhu, sp. nov. — Fig. 19 Proximus L. robinsonii Ridl., sed ramulis costis nervis subtus, pedunculis et calycibus sparsim hirsutis, nervis 5–6-binatis, nervulis subparallelis, petiolis 6–8 mm longis, bracteis late-linearis costatis 1.5 mm longis et lobis calycis lanceolatis nervatis differt. — Typus: Church et al. 2639 (holo L; iso K), Borneo, W Kalimantan, Serawai, 1600 m alt., in upper montane mossy forest, 22 Oct. 1995. Treelet, 2 m high; branchlets terete, c. 1.5 mm diam, deeply brown, sparsely hirsute. Leaves: blades oblong, 6–13 by 2.5– 3.5 cm, chartaceous, glabrous except sparsely hirsute midrib and nerves beneath, apex acuminate to shortly caudate, base acute, midrib slightly depressed above, prominent beneath, nerves flat above, prominent beneath, 5–6 pairs, nervules obscure above, conspicuous beneath, subparallel; petioles 6–8 mm long, hirsute. Stipules lanceolate, c. 3 mm long, hirsute. Inflorescences peduncled; peduncles slender, c. 0.3 mm diam, 2.5–3 cm long, hirsute; bracts 2 at each node, usually unequal, longer ones linear and with a midrib, up to 1.5 cm long, 1 mm broad, sparsely hirsute; shorter ones narrowly linear, hirsute. Flowers pediceled; pedicels 1–3 mm long, hirsute; calyx hirsute, lobes 4, lanceolate, usually nerved, 5 mm long, hirsute external surface, tube c. 2 mm long; corolla not seen. Drupes subglobose, glabrous, 4 obtuse ridges, verrucose on surface, 4–5 mm diam, crowned by lanceolate calyx lobes. Distribution — Indonesia (W Kalimantan). Habitat & Ecology — In upper montane mossy forest. Flowering in October. Note — This species is characterized by its slender peduncles, linear bracts with a midrib and nerved lanceolate calyx 52 Blumea – Volume 57 / 1, 2012 Fig. 19 Lasianthus lancilobus H.Zhu. a. Habit with inflorescences; b. detail of branch; c. detail of inflorescence; d. calyx (all: A.C. Church et al. 2639, holo L). 5 mm 2 cm 2.5 mm 53 H. Zhu et al.: Revision of Malesian Lasianthus lobes. It is closely related to L. robinsonii, but differs from the latter by having sparsely hirsute branches and leaf midrib and nerves beneath, peduncles and calyx; nerves 5–6 pairs, nervules subparallel; petioles 6–8 mm long; bracts broadly linear with a midrib, up to 1.5 cm long; calyx lobes lanceolate, nerved. Lasianthus robinsonii has spreading tomentose branches, leaf nerves beneath and peduncles; nerves 8 pairs, nervules reticulate; petioles short, less than 3 mm long; bracts subulate-linear, less than 1 cm long and without a midrib; calyx lobes subulate. 66. Lasianthus latifolius (Blume ex DC.) Miq. Lasianthus latifolius (Blume ex DC.) Miq. (1857) 320; (1869) 245; Merr. (1921) 577; Bakh.f. (1965) 342. — Mephitidia latifolia Blume ex DC. (1830) 453. — Type: Blume s.n. (lecto L, designated here, barcode L0057522). Paratypes: Blume s.n. (L, barcode L0057523), Blume s.n. (L, barcode L0057524), Blume s.n. (L, barcode L0057525), Blume s.n. (L, barcode L0057526), Java, Salak. Shrubs; branchlets terete, robust, c. 3 mm diam, glabrous except appressed-puberulous nodes. Leaves: blades ellipticoblong, 13–15 by 5–7 cm, coriaceous, glabrous except sparsely hirsute hairs on midrib and nerves beneath, apex cuspidate, base acute or broadly cuneate, midrib slightly depressed above, prominent beneath, nerves slightly prominent above, prominent beneath, c. 7 pairs, ascending at an angle of 60–70°, curved to the margin, nervules prominent on both surfaces, densely anastomosing, parallel; petioles 1–1.5 cm long, appressed-puberulous. Stipules triangular, c. 3 mm long, appressed-puberulous. Cymes sessile or very small tuberculate (c. 1 mm long); bracts absent. Flowers sessile; calyx campanulate, puberulous, tube c. 1.5 mm long, lobes 5, lanceolate-triangular, c. 2.5 mm long; corolla not seen. Drupes globose, red in colour, c. 5 mm diam, glabrous, 12-ridged; pyrenes 6, sulcate on the abaxial face. Distribution — Indonesia (Java). Note — This species is closely related to L. calycinus Dunn from Hainan of China. 67. Lasianthus laxifloroideus H.Zhu, sp. nov. — Fig. 20 Proximus L. laxifloro Merr. sed ramulis costia nervis subtus stipulis pedunculis calycis appressi-pubescentibus, foliis angustioribus, petiolis 5 mm longis, stipulis minoribus triangularis 2 mm longis, cymis 1–2 cm longis differt. A L. dinghoui H.Zhu foliis oblongis vel ovato-oblongis 2.5–4.5 cm latis apice acuminatis vel cuspidatis, lobis calycis lanceolatis 1.5 mm longis differt. — Typus: Ching S.40794 (holo L; iso K, L), Borneo, Sabah, Kelingkang Range, 1650 ft, in mixed dipterocarp forest, 10 Oct. 1979. Shrub, up to 3 m high; branchlets terete, c. 1.5 mm diam, densely appressed-pubescent. Leaves: blades oblong or ovateoblong, 10–16 by 2.5–4.5 cm, subcoriaceous, glabrous except appressed-pubescent midrib and nerves beneath, apex acuminate to cuspidate, base abruptly acute to subrounded, midrib and nerves slightly prominent above, conspicuously prominent beneath, nerves 3–6 pairs, nervules prominent on both surfaces, more conspicuous beneath, parallel; petioles c. 5 mm long, appressed-pubescent. Stipules triangular, c. 2 mm long, appressed-pubescent. Inflorescences peduncled, appressed-pubescent; peduncles 0.5–2 cm long, trifurcate at each node; bracts subulate-linear, 2–3 mm long, pubescent; bracteoles small, subulate, c. 1 mm long. Flowers pediceled; pedicels 3–5 mm long, articulate, appressed-pubescent; calyx campanulate, appressed-puberulous, tube 1.5 mm long, lobes 4, oblong-lanceolate, c. 1.5 mm long; corolla glabrous external surface, 6–7 mm long, lobes 4. Drupes rhomboidal-globose, 5 mm diam, glabrous, verrucose external surface, conspicuously 4-ridged; pyrenes 4, strongly verrucose on abaxial face and sides. Distribution — Malaysia (Borneo). Habitat & Ecology — In lower montane forests between 350– 900 m altitude. Flowering in April; fruiting in October. Additional specimens examined. W Kalimantan, Serawai, A.C. Church et al. 2001 (L); W Kalimantan, Kabupaten Sanggau, W. de Jong 889 (L); Sarawak, Lubok Antu, Dyg Awa et al. 44116 (L); Sabah, Kelingkang Range, Rena et al. S.58966 (L); Sabah, Serian, B. & K. Bremer 1650 (L); Sarawak, G. Lesong, B. Lee S.43203 (L). Note — This species is closely related L. laxiflorus and L. dinghoui H.Zhu. It differs from L. laxiflorus by having appressed-pubescent branches, leaf midrib and nerves beneath, stipules, peduncles, and calyx; petioles 5 mm long; stipules small, triangular, c. 2 mm long; inflorescence 1–2 cm long. While L. laxiflorus is a glabrous plant and has petioles 6–10 mm long, stipules lanceolate 5–7 mm long, inflorescence 3–4 cm long. It differs from L. dinghoui by having densely appressedpubescent on branches, and leave beneath, leaves oblong to ovate-oblong, more than 2.5 cm broad; leaf apex acuminate to cuspidate; calyx lobes oblong-lanceolate, c. 1.5 mm long. 68. Lasianthus laxiflorus Merr. Lasianthus laxiflorus Merr. (1937) 298. — Type: Winkler 430 (holo HBG; iso HBG, NY), W Borneo, Bukit Mulu, 450 m alt. Shrubs to treelets, 3 m high, glabrous; branchlets terete, c. 2.5 mm diam, slightly compressed in the youngest part. Leaves: blades oblong, 10–19 by 2.5–6 cm, coriaceous, apex cuspidate, base acute to broadly cuneate, midrib and nerves slightly depressed above, prominent beneath, nerves 3–5, ascending at an angle of 50–60°, curved to the margin, nervules distinct on both surfaces, subreticulate; petioles 6–12 mm long. Stipules triangular to triangular-lanceolate, up to 5 mm long, glabrous. Cymes pedunculate; main peduncles up to 3 cm long, slender, trifurcate; branches 5–10 mm long; bracts subulate-linear, 2–3 mm long. Flowers subsessile, articulate; calyx obconical, with 4–5 teeth, tube c. 2 mm long, teeth subulate, c. 1 mm long; corolla c. 8 mm long, subglabrous, lobes 4–5, oblong-ovate; anthers 4–5, oblong; ovary 4-locular. Drupes ellipsoid, c. 4 mm diam verrucose; pyrenes 3–4. Distribution — Endemic to Borneo (Sarawak, Sabah, Kalimantan). Note — This species is strongly characterized by being glabrous, having relatively large subcoriaceous oblong leaves (13–19 by 5–6 cm) with acute or cuspidate apex and broadly acute base, midrib and nerves conspicuously prominent beneath, stipules subulate-lanceolate up to 5 mm long, cymes peduncled, usually two peduncles in each axil, main peduncles up to 3 cm long, bracts subulate-linear, 3 mm long. 69. Lasianthus ledermannii Valeton Lasianthus ledermannii Valeton (1927) 108. — Type: Ledermann 9373b (B†), New Guinea, Etappenberg, 850 m; Takeuchi 5096 (neo L, designated here; isoneo K), New Guinea, East Sepik, Hunstein range, 915 m alt., 29 Sept. 1989. Shrubs or treelets, 1.5 m high; branchlets terete, robust, c. 3 mm diam, densely dark-brown-tomentose. Leaves: blades elliptic, elliptic-oblong, 8–13 by 3–5 cm, coriaceous, glabrous above except hirsute hairs on midrib, densely brown-tomentose beneath, apex cuspidate or shortly caudate, margin slightly reflexed, base cuneate to subrounded, rarely asymmetric, midrib and nerves flat to slightly depressed above, prominent beneath, nerves 6–9 pairs, ascending at an angle of 45–60°, curved to the margin, nervules obscure above, distinct and prominent beneath, subparallel; petioles c. 5 mm long, densely tomentose. Stipules triangular-lanceolate, c. 5 mm long, densely tomentose. Cymes sessile; bracts numerous, lanceolate to linear-lanceolate, 5–8 mm long, densely tomentose. Flowers 54 Blumea – Volume 57 / 1, 2012 Fig. 20 Lasianthus laxifloroideus H.Zhu. a. Habit with inflorescences; b. detail of lower surface of leaf showing parallel nervules; c. detail of node showing stipule and axillary inflorescences; d. calyx; e. fruit; f. cross section of fruit (all: Y.P. Ching S.40794, holo and iso L). 1 mm 2 cm 2 mm 55 H. Zhu et al.: Revision of Malesian Lasianthus sessile; calyx obconical, densely tomentose, tube c. 1 mm long, lobes 5, lanceolate, c. 1.5 mm long; corolla tube 4–5 mm long, glabrous, lobes 5, ovate, c. 3 mm long, hairy both sides. Drupes globose to subglobose, 4–5 mm diam, tomentose; pyrenes 5, smooth on the abaxial face. Distribution — Indonesia (Irian Jaya) and Papua New Guinea. Notes — According to Valeton, this species has fuliginoustomentose hairs on branches, leaves nerves beneath, bracts, and drupes; leaves short petiolate, elliptic-oblong or ellipticlanceolate, subcoriaceous; midrib above and nerves beneath tomentose; nerves 6–7; stipules linear, hirsute; bracts numerous, subulate-linear, outer ones 7–10 mm long; calyx hirsute, lobes 2 mm long; drupes globose, apex hirsute. The type of L. ledermannii was destroyed during World War II in Berlin. There are several specimens from New Guinea, which agree well with the description of L. ledermannii. Among them, Takeuchi 5096 perfectly matches the type description. Therefore we select it as a neotype. This species is closely related to the western Malaysian species L. ellipticus. 70. Lasianthus linearifolius H.Zhu, sp. nov. Similis L. angustifolius King & Gamble, sed ramulis et foliis glabris, foliis linearibus, lobis calycis triangularibus 4 differt. — Typus: Beaman et al. 7783 (holo L), Borneo, Sabah, Ranau, 750 m alt., in dipterocarp forest, on ultramafic soil, 8 Dec. 1983. Shrub, 1 m high; branchlets terete, slender, less than 1 mm diam, sparsely puberulous or glabrous. Leaves: blades linear, 10–14 by 0.9–1.8 cm, subcoriaceous, glabrous or except sparsely pubescent midrib beneath, apex long-caudate, margin slightly undulate, base acute, midrib slightly prominent above, conspicuously prominent beneath, nerves slender, slightly prominent above, prominent beneath, 10–16 pairs, ascending at an angle of 80–90°, curved near margin, going paralleling to margin for some distance and joining the nerves above, forming an obscure intramarginal vein, nervules obscure above, weakly prominent beneath, subparallel; petioles 3–6 mm long, sparsely puberulous. Stipules small, subulate, c. 1 mm long, pubescent. Inflorescences sessile; bracts absent. Flowers fascicular, sessile; immature calyx campanulate, pubescent or glabrous, tube c. 1 mm long, lobes 4, triangular, up to 1 mm long; corolla pubescent on external surface. Drupes subglobose, with 4 obtuse ridges, c. 3 mm diam, puberulous or glabrous, crowned by 4 small triangular calyx lobes; pyrenes 4. Distribution — Malaysia (Sabah). Habitat & Ecology — In dipterocarp forest of SE base of Mt Kinabalu on ultramafic soil. Flowering and fruiting in December. Additional specimen examined. Sahab: Ranau, D. Andau 1049 (K). Note — This species is characterized by its linear leaves. It differs from L. angustifolius by having almost glabrous branchlets and leaves, leaves linear, calyx with 4 triangular lobes. Lasianthus angustifolius has branchlets densely pubescent, leaves oblong-lanceolate, leaf midrib and nerves densely pubescent, calyx with 5 triangular teeth. 71. Lasianthus longibracteatus H.Zhu, sp. nov. — Fig. 21 Proximus L. rhinocerotis Blume, sed foliis majoribus 26–40 cm longis subtus hispidis, nervulis reticulatis, bracteis conspicue lonioribus, calycibus fissilobis, lobis calycis lineario-lanceolatis 5–6 mm longis differt. A L. megaphyllo H.Zhu foliis subtus hispidis basi subrotundatis, petiolis longioribus, cymis capitatis brevi-pedunculatis, bracteis numerosis 3 cm longis differt. — Typus: Burley et al. 858 (holo L), Kalimantan, S. Kahayan, 150 m alt., in primary lowland forest, 1 May 1988. Shrub, 1–1.5 m high; branchlets robust, 6–7 mm diam, fistulose, compressed, glabrescent. Leaves: blades obovate-oblong, coriaceous, 26–46 by 10–12 cm, glabrous above, hispid hairs on venation beneath, apex cuspidate to obtuse, base subrounded; midrib depressed above, conspicuously prominent beneath, hispid beneath, nerves depressed above, prominent beneath, 12–15 pairs, ascending at an angle of c. 60°, curved to the margin, hispid beneath, nervules depressed above, prominent beneath, reticulate; petioles c. 1 cm long, hirsute. Stipules triangular-lanceolate, c. 1 cm long, 4–5 mm broad at base. Inflorescences axillary, capitate, very short pedunculate or subsessile; peduncles 1–2 mm long; bracts numerous, linear, up to 3 cm long, densely hirsute. Flowers covered by bracts, sessile; calyx split into 5 linear-lanceolate lobes near to the base lobes 5–6 mm long, 1 mm broad, hirsute external surface and densely hirsute at apex, glabrous internal surface; corolla 12 mm long, subglabrous except the upper part; style filiform, up to 15 mm long; stigma 4–5, filiform. Drupes subglabrous, 6 mm diam, conspicuously 6-ridged, smooth; pyrenes 6, smooth on the abaxial face, ovate-triangular in transverse section. Distribution — Endemic to Borneo (Sarawak, Kalimantan). Habitat & Ecology — In lowland dipterocarp forest. Flowering in May; fruiting in February. Additional specimens examined. W Kalimantan, W. de Jong 760 (L), A.C. Church et al. 1681; Borneo, unknown coll., L barcode L0310812, L0310813; Borneo, H. Hallier 2016, Teysmann 8005; Sarawak, W.M.A. Brooke 9348. Note — This species has the biggest leaves and the longest bracts in the genus. It is close to L. rhinocerotis by its similar capitate inflorescence with short peduncles and numerous linear bracts. It differs from the latter by having much larger leaf blades (26–40 cm long) covered with hispid hairs beneath, and with reticulate nervules; bracts up to 3 cm long; calyx split to the base; calyx lobes linear-lanceolate, 5–6 mm long. It differs from L. megaphyllus by having leaves with subrounded base, hispid beneath; leaf venation more prominent beneath; petioles 1 cm long; inflorescence capitate, very short pedunculate; bracts numerous, linear, up to 3 cm long, densely hirsute. 72. Lasianthus longifolius Wight Key to the subspecies and varieties 1. Leaves with rigid, minute hairs on venation beneath; nervules conspicuously prominent beneath; corolla shorter, less than 1 cm long; drupes crowned by a short cupular calyx limb (not enlarged) ...................... a. subsp. longifolius 1. Leaves glabrous on both surfaces; nervules prominent on both surfaces; corolla longer, up to 2.5 cm long; drupes crowned by an enlarged cupular calyx limb...........2 2. Leaves obovate-oblong to lanceolate-oblong, 6–11 cm broad, apex cuspidate to cuspidate-acute, base cuneate . ................................. b. subsp. laevis 2. Leaves broadly oblong, 11.5–13.5 cm broad, the apex rounded, the base subrounded ...... c. var. obtusifolius a. subsp. longifolius Lasianthus longifolius Wight (1846) 514; Hook.f. (1880) 187; King & Gamble (1904) 116; Ridl. (1923a) 164. — Type: Kew Dist. 2936 (Griffith s.n.) (holo K), Peninsular Malaysia. Treelets, c. 2 m high; branchlets robust, often compressed, c. 3–5 mm diam, glabrous. Leaves: blades obovate-oblong to lanceolate-oblong, 20–33 by 6–11 cm, coriaceous, glabrous above, rigid minute hairs on venation beneath and scabrous (asperate) when touching, apex cuspidate to cuspidate-acute, base cuneate, slightly asymmetric, midrib and nerves slightly 56 Blumea – Volume 57 / 1, 2012 Fig. 21 Lasianthus longibracteatus H.Zhu. a. Habit with inflorescences; b. detail of lower surface of leaf; c. bracts; d. flower; e. calyx; f. calyx split opened; g. fruit; h. cross section of fruit (all: J.S. Burley et al. 858, holo L). 4 mm 2 cm 3 mm 6 mm 5 mm 5 mm 57 H. Zhu et al.: Revision of Malesian Lasianthus Fig. 22 Lasianthus longifolius Wight subsp. laevis H.Zhu. a. Habit with inflorescences; b. lower surface of leaf showing reticulate nervules; c. detail of node showing axillary inflorescences; d. flower; e. calyx; f. fruit (a–e: C. Hansen 295, holo L; f: Yii et al. S.43146, MO). 2 cm 5 mm 5 mm 64 Blumea – Volume 57 / 1, 2012 with an obconical tube and campanulate limb, tube 0.8–1 mm long, glabrous, limb with 4 lanceolate lobes which are 0.6–8 mm long and hirsute; corolla 8–10 mm long, tube glabrous outside, villose inside, lobes 4, hairy on both sides. Drupes oblong, 5 mm diam, puberulous; pyrenes 4–5. Distribution — Peninsular Malaysia, Indonesia (Sumatra). Note — This species is very close to L. chrysoneurus but differs from the latter by having relatively small leaves, stipules and bracts, and denser but less appressed pubescence. 81. Lasianthus mucronulatus (Korth.) Miq. Lasianthus mucronulatus (Korth.) Miq. (1857) 325; (1869) 249. — Mephitidia mucronulata Korth. (1851) 222. —- Type: Korthals s.n. (lecto L, designated here, barcode L0057756), Sumatra, Singalang; Korthals s.n. (para, n.v.), Borneo, Balaran, and Pamaton. Lasianthus palembanicus Miq. (1861) 549. — Lasianthus stercorarius Blume var. palembanicus (Miq.) Miq. (1869) 248. — Type: Teysmann s.n. (holo U), Sumatra, Palembang, Ogan-ulu. Lasianthus stercorarius Blume var. borneensis Miq. ex Valeton & Winkler (1910) 569, syn. nov. — Type: Winkler 2216 (n.v.), Borneo. Shrubs to treelets; branchlets terete, c. 2 mm diam, puberulous, dark-brown when dry. Leaves: blades oblong-lanceolate, 10–12 by 2–4 cm, chartaceous, glabrous or sparsely pubescent hairs on midrib above, pubescent hairs on venation beneath, apex caudate-acuminate or acuminate, margin slightly undulate and reflexed, base broadly cuneate to subrounded, slightly asymmetric, midrib and nerves slightly prominent on both surfaces, nerves slender, 10–16 pairs, ascending at an angle of 45–60°, curved to the margin; nervules distinct on both surfaces, reticulate; petioles 2–4 mm long, puberulous. Stipules triangular, usually 2 mm long, puberulous. Cymes sessile; bracts absent. Flowers sessile; obconic-campanulate; calyx tube c. 0.8 mm long, glabrous or pubescent, calyx limb c. 0.6 mm long, with 4–5 minute and pubescent teeth; corolla pubescent outside, tube c. 2 mm long, lobes c. 1 mm long. Drupes globose, c. 3 mm diam, puberulous or subglabrous; pyrenes 5. Distribution — Indonesia (Sumatra, Java, Kalimantan). Notes — This species differs from L. stercorarius by having usually relatively large chartaceous leaves with broadly cuneate to subrounded bases and more than 10 paired nerves and small, 2 mm long, triangular stipules and smaller drupes without a truncate calyx limb. Lasianthus mucronulatus var. lineata was established on Korthals’s Sumatra specimens. It has triangular stipules with swollen base, subcoriaceous leaves with much acute base which are the same as L. stercorarius. Var. lineata here is reduced to L. stercorarius. Lasianthus stercorarius var. borneensis is the same as L. mucronulatus morphologically as pointed by Valeton & Winkler. It is here reduced to a synonym of L. mucronulatus. 82. Lasianthus myrtifolius Ridl. Key to the subspecies 1. Leaves ovate-lanceolate or ovate with a caudate-acuminate apex; calyx and corolla pubescent .. a. subsp. myrtifolius 1. Leaves oblong with an abruptly acute apex; calyx and corolla glabrous ......................... b. subsp. vicarius a. subsp. myrtifolius Lasianthus myrtifolius Ridl. (1909) 35; (1923a) 161; K.M.Wong (1989) 369. — Type: Ridley 13630 (holo SING; iso K), Peninsular Malaysia, Pahang, 5000 ft. Shrubs, 1–2 m high; branchlets terete, slender, c. 1 mm diam, puberulous. Leaves: blades ovate-lanceolate to ovate, 3–7 by 1–2 cm, coriaceous, glabrous and nitid on both surfaces, or minutely puberulous hairs on midrib and nerves beneath, apex caudate-acuminate, aristate, arista c. 1 mm long, base acute, midrib slightly depressed above, prominent beneath; nerves prominent on both surfaces, 5–6 pairs, ascending at an angle of over 45°, curved to the margin, nervules prominent and distinct on both surfaces, parallel; petioles 3–5 mm long, puberulous. Stipules subulate-triangular, c. 1 mm long, pubescent. Cymes sessile; bracts absent. Flowers sessile; calyx campanulate, c. 2 mm long, pubescent outside, lobes 4, triangular, c. 1 mm long; corolla c. 5 mm long, lobes 4, linear, pubescent outside, hairy inside. Drupes globose, c. 4 mm diam, verrucose, conspicuously 4-ridged; pyrenes 4, sulcate on the abaxial face and sides. Distribution — Endemic to Peninsular Malaysia. Note — This species has small thickly coriaceous leaves, which look like myrtaceous leaves, and very small flowers. b. subsp. vicarius H.Zhu, subsp. nov. — Fig. 26 A typo foliis majoribus oblongis apice abrupte acutis, calycibus et corollis extus glabris differt. Similis L. coriaceifolius H.Zhu, sed nervulis foliorum parallelis, calycibus minoribus, lobis calycis triangularibus, drupis non conspicue porcatis, pyrenis abaxiali laevigatis differt. — Typus: Banying ak Nyudong S.19397 (holo L), Borneo, Naoung, Anap, 1150 m alt., in submontane heath forest, 11 Oct. 1963. Treelet, 5–6 m high; branchlets terete, robust, c. 2.5 mm diam, glabrous, darkish. Leaves: blades oblong, 4.5–7.5 by 2–3 cm, thickly coriaceous, glabrous, apex abruptly acute, margin recurved, base acute, midrib, nerves and nervules prominent on both surfaces, nerves 6–8 pairs, nervules parallel; petioles c. 3 mm long, glabrous, darkish. Stipules triangular-subulate, 1.5 mm long, subglabrous. Inflorescences sessile; bracts small, ovate, 1 mm long or inconspicuous. Flowers sessile; calyx campanulate, 2–3 mm long, glabrous, darkish, lobes 4, triangular, c. 1 mm long; corolla 4–5 mm long, glabrous on external surface, darkish, lobes ovate-triangular, 1 mm long. Drupes subglobose, c. 4 mm diam, glabrous, crowned by calyx lobes; pyrenes 4, smooth on abaxial face, suborbicular in the transverse section. Distribution — Borneo (Sarawak). Habitat & Ecology — Submontane heath forest or mossy forest on the summit over 1100 m altitude. Flowering and fruiting between May and October. Additional specimen examined. Sarawak, Bantulu, E.F. Brunig S.8770 (K, L). Note — This subspecies differs from the typical subspecies by having oblong leaves with abruptly acute apex, glabrous calyx and corolla external surface. The typical subspecies has ovate-lanceolate or ovate leaves with caudate-acuminate apex, pubescent calyx and corolla external surface. It differs from L. coriaceifolius by having leaves with parallel nervules, calyx relatively small (2–3 mm long), calyx lobes triangular, drupes without conspicuous ridges, pyrenes smooth on abaxial face. Lasianthus coriaceifolius has leaves with conspicuously reticulate nervules, calyx 5 mm long, calyx lobes ovate-oblong, drupes with 8 acute ridges, pyrenes with ridges on the abaxial face. 83. Lasianthus nervosus King & Gamble Lasianthus nervosus King & Gamble (1904) 129; Ridl. (1923a) 163; K.M.Wong (1989) 367. — Type: Scortechini 342 (holo K; iso BM), Peninsular Malaysia, Perak. Shrubs to treelets, 1–2 m high; branchlets terete, 1.5–2 mm diam, densely tomentose. Leaves: blades oblong to oblong- 65 H. Zhu et al.: Revision of Malesian Lasianthus Fig. 26 Lasianthus myrtifolius Ridl. subsp. vicarius H.Zhu. a. Habit with inflorescences; b. flower (all: Banying ak Nyudong S.19397, holo L). 2 cm 3 mm 66 Blumea – Volume 57 / 1, 2012 lanceolate, 2.5–5 by 1–2 cm, coriaceous, glabrous and nitid above, densely tomentose beneath, especially on venation, apex acute, mucronate, margin reflexed, base broadly cuneate to subrounded, midrib depressed above, strongly prominent beneath, nerves flat above, strongly prominent beneath, 6–8 pairs, ascending at an angle of over 50°, curved to the margin, nervules distinct above, prominent beneath, parallel; petioles 3–5 mm long, densely tomentose. Stipules subulate, c. 1 mm long, densely tomentose. Cymes sessile; bracts absent. Flowers sessile; calyx campanulate, c. 2 mm long, densely tomentose, lobes 4, triangular, c. 0.6 mm long; corolla c. 4 mm long, tomentose outside, lobes 4. Drupes globose, sparsely hairy, c. 4 mm diam; pyrenes 4. Distribution — Peninsular Malaysia, Indonesia (Sumatra, Kalimantan). Notes — This species has densely tomentose hairs on branchlets and leaf midrib and nerves beneath; leaves oblong, 2.5–5 by 1–2 cm with acute and mucronate apex and abruptly acute base, midrib depressed and nerves flat above, midrib and nerves strongly prominent beneath; cymes sessile, absent; flowers 4-merous; calyx tomentose with 4 triangular teeth; drupes globose with 4 pyrenes. This species differs from L. myrtifolius by having densely tomentose hairs on branchlets and leaf midrib and nerves beneath; leaves oblong, nerves flat above; dupes globose without conspicuous ridges. Lasianthus myrtifolius has branchlets thinly puberulous or subglabrous; leaves ovate-lanceolate, thinly puberulous or glabrous beneath, with caudate-acuminate apex; nerves conspicuously prominent above; dupes with conspicuous 4 ridges. 84. Lasianthus nigrescens H.Zhu, sp. nov. Proximus L. sabahensis H.Zhu, sed foliorum laminis oblongis apice cuspidatis basi subrotundatis, nervulis conspicue prominentibus, petiolis 5–7 mm longis et tubis calycis 3 mm longis differt. — Typus: Beaman 8149 (holo K), Malaysia (Sabah), Ranau District, East Mesilau River, 1950–2100 m alt., in oak-laurel forest, 30 Dec. 1983. Shrubs, up to 2.5 m high; branchlets terete, c. 2 mm diam, glabrous, more or less compressed in the youngest part. Leaves: blades oblong, 6–12 by 3–5 cm, coriaceous, glabrous, turning black when dry, apex cuspidate, rarely acute, margin reflexed, base subrounded, rarely acute, midrib and nerves prominent on both surfaces, nerves 6–7 pairs, ascending at an angle of 80–90°, curved to the margin, nervules loosely anastomosing, subparallel, conspicuously prominent on both surfaces; petioles 5–7 mm long, glabrous. Stipules triangular, c. 2 mm long, glabrous. Cymes sessile; bracts absent. Flowers sessile, fascicular; calyx campanulate, tube c. 3 mm long, with 4 c. 1 mm long minute teeth, puberulous or glabrous outside; corolla tubular, c. 1 cm long, puberulous or glabrous outside, lobes 4, c. 2 mm long, puberulous or glabrous. Drupes ellipsoid, 6–7 mm long, c. 5 mm diam, glabrous, crowned by a cupular calyx limb with 4 minute teeth, 2-pyrene, with cork-like structure between pyrenes; pyrenes with a hard wall, smooth on the abaxial face. Distribution — Endemic to Borneo. Habitat & Ecology — In forest between 1800–2200 m altitude. Flowering and fruiting from March to December. Note — Lasianthus nigrescens is close to L. sabahensis, especially in the same type of drupes that have a cork-like structure between pyrenes. Lasianthus nigrescens differs from the latter in having leaves oblong, with usually cuspidate apex and subrounded base, nervules conspicuously prominent on both surfaces; petioles 5–7 mm long; calyx tube 3 mm long, not expended at limb. Lasianthus nigrescens differs from L. membranaceus in having coriaceous leaves with nervules conspicuously prominent on both surfaces; drupes with cork-like structure between pyrenes; pyrene walls smooth on the abaxial face. 85. Lasianthus obliquinervis Merr. Key to the subspecies 1. Leaves with acute base; stipules triangular to oblong-triangular........................ a. subsp. obliquinervis 1. Leaves with cuneate base; stipules linear-subulate ..... ......................... b. subsp. novoguineensis a. subsp. obliquinervis Lasianthus obliquinervis Merr. (1906) 136. — Type: Whitford 247 (lecto US, designated here), Philippines, Luzon, Bataan, May 1904. Shrubs, c. 2 m high, branches 5–6 mm diam, striate; branchlets 2–3 mm diam, terete or slightly compressed, puberulous. Leaves: blades ovate-lanceolate to ovate-elliptic, 10–15 by 2.5–5 cm, subcoriaceous, glabrous above, puberulous beneath, brownish, apex acuminate to acute, rarely cuspidate, base acute, slightly asymmetric and decurrent, midrib slightly depressed above, prominent beneath, pubescent beneath, nerves flat above, prominent beneath, 3–6 pairs, ascending at an angle of 50–70°, curved to the margin, brown pubescent, nervules obscure above, conspicuous beneath, parallel, glabrous; petioles 1.5–2 cm long, pubescent. Stipules triangular to oblong-triangular, up to 5–6 mm long, 3–4 mm broad at base, pubescent. Inflorescences sessile; bracts small, ovate to subulate, c. 1 mm long, hirsute. Flowers fascicular, sessile; calyx campanulate, 1.5 by 1.5 mm, densely pubescent, limb with 4–5 triangular lobes, lobes 0.5 mm long, densely hirsute outside, glabrous inside; corolla 6–8 mm long, densely hirsute except the base, lobes 5, oblong, c. 4 mm long; stamens 5, filaments 2 mm long; anthers 1 mm long. Drupes globose, hirsute, blue when fresh, 4–5 mm diam, crowned by persistent calyx lobes, 5–8-ridged, often 6-ridged; pyrenes 5–8, smooth on the abaxial face, triangular in transverse section. Distribution — Philippines (Luzon to Mindanao), Papua New Guinea (Southern Highlands). Habitat & Ecology — Montane forest between 800–1800 m altitude. Notes — This species is close related to L. chrysoneurus. It differs from the latter by having leaves with longer petioles (1.5–2 cm) and acute base, as well as less nerves (3–6 pairs); bracts small, less than 2 mm long; calyx lobes triangular, 1.5 mm long, glabrous internal surface; drupes conspicuously 5–8-ridged. Lasianthus chrysoneurus has relatively big ovateoblong or oblong leaves with petioles less than 1 cm long, the base cuneate to obtuse, nerves 8–10 pairs; bracts longer than 2 mm; calyx lobes lanceolate, longer than 1.5 mm, hirsute both surfaces; drupes without conspicuous 5–8 ridges. Lasianthus obliquinervis is variable in leaf size. Specimens which were treated as L. obliquinervis were mixed with L. verticillatus even if identified by Merrill himself. From our checking on specimens in L, MO, SING and Chinese herbaria, most specimens indicated as L. obliquinervis agree well with L. verticillatus, while some do not. When Merrill described L. obliquinervis, he cited 10 collections (syntypes). We are not sure if all the syntypes represent the same species. Only two syntypes, i.e. Whitfort 247 and Meyer 2212 were traced and seen. Both of them match Merrill’s description and surely represent L. obliquinervis. Lasianthus obliquinervis differs from L. verticillatus by having stipules oblong-ovate, c. 5 mm long, calyx limbs with 4–5 triangular lobes and drupes with usually 6 triquetrous pyrenes and not crowned by an enlarged cupular and truncate calyx limb. 67 H. Zhu et al.: Revision of Malesian Lasianthus Fig. 27 Lasianthus obliquinervis Merr. subsp. novoguineensis H.Zhu. a. Habit with inflorescences; b. detail of node showing stipule and axillary inflorescences; c. calyx split opened; d. corolla split opened; e. fruit; f. cross section of fruit (all: J. Vandenberg et al. NGF.39936, holo L). 3 mm 2 cm 2 mm 0.5 mm 68 Blumea – Volume 57 / 1, 2012 b. subsp. novoguineensis H.Zhu, subsp. nov. — Fig. 27 A typo foliis basi asymmetrice cuneatis, stipulis lineari-subulatis 6 mm longis differt. — Typus: J. Vandenberg et al. NGF.39936 (holo L), New Guinea, Western Highlands District, 4450 ft alt., in Castanopsis forest, 30 Oct. 1968. Shrub, branchlets terete, densely puberulous. Leaves: blades lanceolate to elliptic-lanceolate, 10–15 by 2.5–5 cm, subcoriaceous, glabrous above, puberulous beneath, apex acuminate, base cuneate and asymmetric, nerves flat above, prominent beneath, 6–7 pairs, pubescent, nervules conspicuous beneath, parallel. Stipules linear-subulate, 6 mm long, pubescent. Flowers fascicular, sessile; calyx campanulate, densely pubescent, limb with 4–5 triangular lobes; corolla 6–8 mm long, densely hirsute except the base. Drupes globose, hirsute, 4–5 mm diam, crowned by persistent calyx lobes, with 6 obtuse ridges; pyrenes 6, smooth on the abaxial face, ovate in transverse section. Distribution — Endemic to Papua New Guinea (Western Highlands). Habitat & Ecology — In montane forest between 1400–1500 m altitude. Flowering and fruiting between July and October. Additional specimen examined. New Guinea, Western Highlands, Streimann 8460 (L). Note — This subspecies differs from the typical subspecies by having leaves with cuneate and asymmetric bases, stipules linear-subulate (6 mm long). The typical subspecies has leaves with acute and symmetrical bases, stipules triangular to oblongtriangular. 86. Lasianthus oblongatus Merr. Lasianthus oblongatus Merr. (1937) 299. — Type: Winkler 1364 (iso NY), W Borneo, Sungei Obat, 100 m alt. Shrubs; branchlets terete, c. 2 mm diam, slightly appressedpubescent. Leaves: blades narrowly oblong, 9–14 by 2–3 cm, chartaceous, glabrous above, appressed-pubescent hairs on midrib and nerves beneath, apex acuminate, base broadly cuneate to subrounded, midrib and nerves slightly prominent on both surfaces, nerves slender, 13–15 pairs, ascending at an angle of c. 60°, curved to the margin, nervules slightly obscure on both surfaces, subparallel; petioles c. 2 mm long, pubescent. Stipules lanceolate, 4–5 mm long, hirsute, deciduous. Cymes sessile; bracts with a midrib, leaf-like, oblong-lanceolate, c. 1 cm long and 2 mm broad, paired in leaf axil, 4 in each node. Flowers solitary, sessile. Drupes ovoid, c. 4 mm diam, glabrous or sparsely pubescent, calyx lobes persistent, 1.5–2 mm long, hirsute; pyrenes 5. Distribution — Endemic to Borneo. Note — This species is distinct by rather thin oblong leaves and pairs of persistent lanceolate bracts. 87. Lasianthus oblongus King & Gamble Lasianthus oblongus King & Gamble (1904) 127; Ridl. (1923a) 166; K.M.Wong (1989) 368. — Type: King’s collector 4128 (lecto SING, selected by Wong 1989; isolecto BM, K, L). Paratypes: Scortechini 265 (n.v.); Ridley 4935 (n.v.), 9702 (n.v.), 9729 (n.v.); Wray 2590 (SING); King’s collector 402 (SING), 4462 (K), 10082 (n.v.); Curtis 2081 (SING), Perak. Ridley 7438 (n.v.), Selangor, Peninsular Malaysia. Shrubs, up to 2 m high; branchlets terete, c. 2 mm diam, minutely puberulous or glabrescent, dark when dry. Leaves: blades narrowly oblong-lanceolate, 9–14 by 2–3 cm, thinly coriaceous, glabrous above, puberulous hairs on midrib and nerves beneath, apex caudate-acuminate, base acute, dark when dry, midrib and nerves slightly prominent above, prominent beneath, nerves 6–7 pairs, ascending at an angle of 50–60°, curved to the margin; nervules slender, densely anastomosing, parallel; petioles 5–8 mm long, puberulous. Stipules triangular, c. 3 mm long, densely pubescent. Cymes sessile, bracts absent. Flowers pediceled; pedicels c. 3 mm long, glabrous; calyx shallow cupular, c. 1.5 mm long, glabrous or puberulous, limb c. 1 mm long, truncate; corolla c. 5 mm long, pubescent outside, lobes 5, oblong, densely long-floccose inside. Drupes globose, c. 3–4 mm diam, puberulous or glabrous, verrucose, with 7–8 obtuse ridges; pyrenes 7–8. Distribution — Endemic to Peninsular Malaysia. Note — Lasianthus oblongus has oblong-lanceolate leaves with clearly parallel nervules, pedicels relatively short, 2–3 mm long, drupes with 7–8 pyrenes, differing from L. purpureus. 88. Lasianthus obovatibracteatus H.Zhu, sp. nov. — Fig. 28 Proximus L. inodoro Blume subsp. pubescenti H.Zhu, sed foliis basi asymmetrice rotundatis, stipulis minoribus, bracteis spathulato-obovatis minoribus, lobis calycis 4 et drupis glabris differt. A L. sterophyllo Merr. foliis minoribus basi asymmetrice rotundatis, stipulis minoribus, bracteis spathulato-obovatis et pyrenis 3–4 differt. — Typus: Beaman 10038 (holo L; iso K), Borneo, Sabah, Lahad Datu, 800–880 m alt., in low mossy primary forest, on ultramafic rock soil, 10 June 1984. Treelet, c. 2.5 cm high; branchlets terete, c. 2 mm diam, puberulous, darkish. Leaves: blades ovate-oblong, 5–8 by 2–3 cm, thickly coriaceous, glabrous except puberulous midrib beneath, darkish, apex acute, margin slightly recurved, base rounded or subrounded, slightly asymmetric, midrib slightly depressed above, prominent beneath, nerves prominent on both surfaces, 6–7 pairs, ascending at an angle of 70–80°, curved to the margin, nervules prominent on both surfaces, subreticulate; petioles 5–6 mm long, puberulous. Stipules triangular, 3 mm long, subglabrous. Inflorescences sessile; bracts 2, spatulate-obovate, 3 by 2 mm, puberulous or subglabrous. Flowers in fascicle, sessile; calyx campanulate, sparsely puberulous, 4–5 mm long, tubes obconical, 2–2.5 mm long, lobes 4–5, ovate-triangular, c. 2.5 mm long; corolla c. 8 mm long, glabrous outside, lobes 4, ovate-oblong, c. 3.5 mm long, 2.5 mm broad. Drupes ellipsoid, compressed, c. 5 mm diam, glabrous, crowned by 4–5 ovate-triangular calyx lobes; pyrenes 3–4, smooth on the abaxial face. Distribution — Malaysia (Sabah). Habitat & Ecology — Low mossy primary forest on ultramafic rock soil. Flowering and fruiting between March and June. Note — This species is closely related to L. inodorus subsp. pubescens, but differs from the latter by having leaves with rounded and slightly asymmetric base, relatively small stipules, bracts relatively small and spatulate-obovate, drupes glabrous with 3–4 pyrenes. Lasianthus inodorus subsp. pubescens has leaves with cuneate or acute base, stipules 4 mm long, densely pubescent, bracts orbicular and densely pubescent, up to 5 mm diam, calyx lobes 5, drupes pubescent. Lasianthus obovatibracteatus is also similar to L. sterrophyllus, but differs from the latter by having relatively small leaves with rounded and slightly asymmetric base, relatively small stipules, bracts spatulate-obovate, drupes with 3–4 pyrenes. Lasianthus sterrophyllus has leaves 9–12 cm long with acute to subacute base, bracts absent, stipules 4–5 mm long, drupes with 5 pyrenes. 89. Lasianthus obscurus (Blume ex DC.) Miq. — Map 3 Lasianthus obscurus (Blume ex DC.) Miq. (1857) 317; (1869) 245, p.p. excl. specimen Zollinger 621; Hook.f. (1880) 183; Bakh.f. (1965) 341; H.Zhu (2002) 80. — Mephitidia obscura Blume ex DC. (1830) 453. — Type: Blume s.n. (holo L, barcode L0057529), Java. Lasianthus kurzii Hook.f. var. sylvicola H.S.Lo (1993) 3; H.Zhu (1998) 153; H.S.Lo (1999) 90. — Type: E. Hainan Expedit. 530 (holo SCBI), China, Hainan. Lasianthus kurzii Hook.f. var. fulvus C.Y.Wu & H.Zhu (in Zhu 1994) 60. — Type: Yunnan-expedit. 59-13346 (holo KUN), China, Yunnan, Xishuangbanna. 69 H. Zhu et al.: Revision of Malesian Lasianthus Fig. 28 Lasianthus obovatibracteatus H.Zhu. a. Habit with inflorescences; b. detail of lower surface of leaf showing subreticulate nervules; c. detail of node showing stipule and axillary inflorescences; d. calyx; e. fruit; f. cross section of fruit (all: J.H. Beaman 10038, holo L). 1 mm 2 cm 3 mm 70 Blumea – Volume 57 / 1, 2012 Fig. 29 Lasianthus palawanensis H.Zhu. a. Habit with inflorescences; b. detail of lower surface of leaf showing reticulate nervules; c. detail of node showing stipule and axillary inflorescences; d. calyx; e. calyx split opened; f. stipule (all: A.D.E. Elmer 13252, holo L). 3 mm 2 cm 2 mm 71 H. Zhu et al.: Revision of Malesian Lasianthus Shrubs, to 3 m high; branchlets terete, robust, c. 3 mm diam, densely spreading tomentose. Leaves: blades elliptic-oblong to oblong, 12–20 by 3–5 cm, membranaceous, or thinly coriaceous, glabrous above except sometimes tomentose hairs on midrib, tomentose beneath, apex acuminate to cuspidate-acute, base cuneate to subrounded, midrib and nerves slightly prominent above, prominent beneath, nerves 7–10 pairs, ascending at an angle of 50–60°, curved to the margin, nervules distinct on both surfaces, parallel; petioles 5–10 mm long, spreading tomentose. Stipules triangular or oblong-lanceolate, up to 8 mm long, tomentose. Cymes sessile; bracts persistent, outer bracts ovate, inner bracts lanceolate, all not more than 5 mm long, densely tomentose. Flowers sessile or subsessile, numerous; calyx campanulate, 2–3 mm long, tomentose, lobes triangular, 4–6, c. 1–1.5 mm long; corolla 5–6 mm long, puberulous outside in the upper half, pubescent inside at the throat. Drupes subglobose, 4–6 mm diam, tomentose, crowned by persistent calyx lobes; pyrenes 5–6. Distribution — China (Yunnan, Hainan), Vietnam, Thailand, Indonesia (Java, Sulawesi). Notes — Lasianthus obscurus is very close to L. chrysoneurus. Lasianthus obscurus has densely spreading tomentose hairs on branches, leaf undersides, stipules, bracts, and calyx, while L. chrysoneurus has appressed pubescence on branches, leaf undersides, stipules, bracts, and calyx. They are separated only by different indumentum. Considering that there are relatively few diagnostic characters in the genus and the clear distinction between the two taxa by different types of indumentum, it seems better to treat them as two separate species. Lasianthus kurzii var. sylvicola and var. fulvus from mainland SE Asia are conspecific with L. obscurus, and L. kurzii var. kurzii from India and mainland SE Asia is conspecific with L. chrysoneurus. 90. Lasianthus palawanensis H.Zhu, sp. nov. — Fig. 29 Proximus L. trichophlebus Hemsl., sed foliis obovato-oblongis chartaceis, nervulis subreticulatis, lobis calycis 5 subulato-lanceolatis aequalis tubo differt. — Typus: Elmer 13252 (holo L), Philippines, Palawan Island, May 1911. Shrub; branchlets terete, robust, c. 2.5–3 mm diam, sparsely hirsute, densely at nodes, dark-brownish. Leaves: blades obovate-oblong, 10–12 by 3–3.5 cm, chartaceous, glabrous and dark-brownish above, puberulous or glabrescent and lightly brownish beneath, apex cuspidate or shortly caudate, mucronate, base acute; midrib flat above, prominent and sparsely hirsute beneath, nerves slightly prominent above, prominent and hirsute beneath, c. 11 pairs, ascending at an angle of c. 60°, curved gradually to margin, nervules slightly prominent on both surfaces, obscure beneath, reticulate; petioles 3–5 mm long, hirsute. Stipules triangular-lanceolate, 6–7 mm long, hirsute. Inflorescences sessile, bracts fewer, linear, c. 2 mm long, hirsute. Flowers fascicular, sessile; calyx campanulate, c. 4 mm long, hirsute, lobes 5, subulate-lanceolate, c. 2 mm long; corolla c. 5–6 mm long, tube glabrous outside, lobes 5, lanceolate, c. 2 mm long, puberulous external surface; ovary with 5 locules. Drupes not seen. Distribution — Endemic to Palawan Islands of Philippines. Note — This species has hirsute hairs on nodes of branchlets, leaf nerves beneath, stipules, and calyx; stipules triangular-lanceolate, 6–7 mm long. It seems closely related to L. trichophlebus, but differs from the latter by having obovateoblong and chartaceous leaves with subreticulate nervules, calyx lobes 5, subulate-lanceolate, as long as calyx tube. Lasianthus trichophlebus has oblong and coriaceous leaves with parallel nervules, calyx lobes 4, triangular, shorter than calyx tube. 91. Lasianthus papuanus Wernham Lasianthus papuanus Wernham (1916) 78; Valeton (1927) 109; Merr. & L.M.Perry (1946) 219. — Type: Kloss s.n. (holo BM), New Guinea, 500 ft. Shrubs; branchlets terete, densely appressed flavi-hirsute. Leaves: blades oblong-lanceolate, 7–11 by 2–3 cm, glabrous above except hairy midrib, appressed-pubescent hairs on venation beneath, apex acute, base broadly cuneate to subrounded, nerves slightly obscure above, prominent beneath, 7–8 pairs, nervules obscure above, subreticulate; petioles 4–5 mm long, densely flavi-pubescent. Stipules subulate, 8 mm long, appressed-pubescent, deciduous. Cymes sessile; bracts absent. Flowers solitary or paired, 5-merous; calyx campanulate, lobes triangular-lanceolate, 2.5 mm long, glabrous inside and pilose outside; corolla tube 4.8 mm long, sparsely hirsute, lobes 5, oblong, c. 1.5 mm long, hirsute outside, barbate inside. Fruits not seen. Distribution — New Guinea. Note — This species is very close to L. chrysotrichus, but differs from the latter by having appressed-pubescence on branches, nerves beneath, stipules up to 8 mm long, 5-merous flowers without bracts, and triangular-lanceolate calyx lobes, while L. chrysotrichus has spreading harsh-hirsute hairs, stipules 2–3 mm long, usually 4-merous flowers with linear bracts and lanceolate calyx lobes. It differs from L. hexandrus by having stipules up to 8 mm long, calyx campanulate, lobes triangular-lanceolate. 92. Lasianthus pedicellatus H.Zhu, sp. nov. — Fig. 30 Proximus L. oblongo King & Gamble, sed stipulis minoribus, calycibus triangulatis dentatis, drupis levigatis obtuse 4–5-porcatis, pyrenis 4–5 differt. — Typus: De Vogel 5344 (holo L), Sulawesi, Palu, Mt Poroka Timbu, 1950 m alt., in primary montane forest, 14 May 1979. Treelet, 2 m high; branchlets terete, c. 1.5 mm diam, dark in the youngest part, glabrous or thinly puberulous. Leaves: blades lanceolate, 6–9 by 1.5–2.5 cm, glabrous except sparsely puberulous midrib and nerves beneath, apex acuminate to caudate, aristate, arista c. 1 mm long, margin recurved, base acute, midrib slightly depressed above, prominent beneath, nerves flat above, prominent beneath, 5 pairs, ascending at an angle of c. 60°; nervules prominent on both surfaces, parallel; petioles c. 5 mm long, sparsely puberulous or glabrescent. Stipules very small, triangular, less than 1 mm long. Inflorescences extremely congested to nearly sessile; bracts absent. Flowers fascicular, pediceled; pedicels slender, c. 2 mm long; calyx obconical, c. 1 mm long, with 5 triangular teeth, puberulous; corolla c. 4 mm long, puberulous outside. Drupes subglobose, c. 3 mm diam, smooth, subglabrous, with 4–5 obtuse ridges; pyrenes 4–5, smooth on the abaxial face, orbicular at the transverse section. Distribution — Malaysia (Sabah) and Indonesia (Sulawesi). Habitat & Ecology — In montane forest between 1500–2000 m altitude. Flowering and fruiting between March and May. Additional specimens examined. Sulawesi, Mt Roroka Timbu, E.F. de Vogel 5136 (K, L); Borneo, Sabah, J.H. Beaman 8928 (K, L); Sabah, Kinabalu, J & M.S. Clemens 29838 (L). Note — This species is closely related to L. oblongus, but differs from the latter by having very small stipules, calyx with triangular teeth, drupes smooth, with 4–5 obtuse ridges and crowned by small triangular calyx teeth, pyrenes 4–5. Lasianthus oblongus has relatively big stipules 3–4 mm long, calyx shallow cupular with truncate limb, drupes verrucose, with 7–8 obtuse ridges and crowned by a short cupular calyx limb, pyrenes 7–8. 72 Blumea – Volume 57 / 1, 2012 Fig. 30 Lasianthus pedicellatus H.Zhu. a. Habit; b. detail of node showing axillary inflorescences; c. detail of node showing fruits; d. flower; e. fruit; f. cross section of fruit (all: E.F. de Vogel 5344, holo L). 1 mm 2 cm 1 mm 1 mm 73 H. Zhu et al.: Revision of Malesian Lasianthus 93. Lasianthus pendulus Ridl. Lasianthus pendulus Ridl. (1909) 37; (1923a) 168; K.M.Wong (1989) 372. — Lasianthus capitatus Blume subsp. pendulus (Ridl.) H.Zhu (2000b) 398. — Type: Ridley 14007 (holo SING; iso K), Peninsular Malaysia, Pahang, Telem. Treelets, c. 2 m high; branchlets terete, robust, c. 3 mm diam, densely spreading long-hairy. Leaves: blades ovate-lanceolate, 7–11 by 2–3.5 cm, coriaceous, glabrous above, spreading longhairy beneath, apex acuminate, margin conspicuously reflexed, base rounded to subcordate, midrib and nerves conspicuously depressed above, strongly prominent beneath, nerves 8–9 pairs, ascending at an angle of over 50°, curved to the margin, nervules slightly obscure, distinct beneath, subparallel; petioles very short, c. 1 mm long, densely long-hairy. Stipules inconspicuous. Cymes pedunculate; peduncles 2–3.5 cm long, densely spreading long-hairy; bracts filiform, numerous, up to 1.5 cm long, densely long-hairy. Flowers sessile; calyx with 5 long, lanceolate-subulate and hairy lobes; corolla glabrous. Drupes obovoid, narrowed to base, 5-ridged, crowned by hairy calyx lobes; pyrenes 5. Distribution — Peninsular Malaysia. Habitat & Ecology — Montane forest, 1500–1900 m altitude. Note — Lasianthus pendulus differs from L. capitatus in having ovate to ovate-lanceolate leaves with rounded base, shorter petioles, nerves conspicuously depressed above and more slender peduncles. 94. Lasianthus perakensis King & Gamble Key to the varieties 1. Leaves with nerves 7–8 pairs, 15–25 by 5.5–8 cm; rustypuberulous on branches, midrib and nerves beneath; drupes c. 4 mm diam, sessile ..............a. var. perakensis 1. Leaves smaller with nerves 3–5 pairs, 13–16 by 3–4.5 cm; sparse puberulous on branches, midrib and nerves beneath; drupes smaller c. 2 mm diam, with 1 mm long pedicels .. ............................b. var. kalimantanensis a. var. perakensis Lasianthus perakensis King & Gamble (1904) 126; Ridl. (1923a) 160; K.M. Wong (1989) 372 (excl. syn. L. flavinervius Ridl.); H.Zhu (2001b) 139. — Type: Wray 4128 (lecto SING, designated by Wong 1989). Paratypes: King’s collector 682 (n.v.), 2438 (n.v.), 2838 (n.v.), 10210 (n.v.), Peninsular Malaysia, Perak. Shrubs; branches terete, robust, c. 3 mm diam, or slightly compressed in the youngest part, minutely rusty-puberulous. Leaves: blades elliptic or obovate-elliptic, 15–25 by 5.5–8 cm, subcoriaceous, glabrous above, puberulous on midrib and nerves beneath, apex cuspidate or caudate, base cuneate; midrib slightly depressed above, prominent beneath, nerves 7–8 pairs, flat above, prominent beneath, nervules subparallel, conspicuous on both sides; petioles about 1 cm long, rusty-puberulous. Stipules triangular, acute, up to 5 mm long, rusty-puberulous. Cymes sessile, bracts absent. Flowers sessile, crowned; calyx campanulate, c. 2 mm long, much tapered to the base, mouth wide, truncate, with 5 indistinct blunt spreading segments, pubescent; corolla short, 3 mm long, lobes ovate-triangular, 1.5 mm long, pubescent; ovary 5-celled. Drupes hemispheric, c. 4 mm diam, subglabrous or sparsely puberulous, 10-ribbed, crowned by the calyx limb; pyrenes 10. Distribution — S Thailand, Malaysia (Peninsular, Sabah) and Indonesia (Sumatra, Kalimantan). Note — Lasianthus perakensis has a wide to slightly trumpetshaped calyx limb with 5 indistinct blunt spreading lobes, drupes hemispheric with 10 obtuse ridges and crowned by blunt calyx lobes, pyrenes 5. b. var. kalimantanensis H.Zhu, var. nov. A typo foliis minoribus, nervis 3–5, ramis sparsim puberscentibus, costis et nervis sparsim puberscentibus subter, pedicellis 1 mm longis, drupis minoribus differt. — Typus: Church et al. 713 (holo K), Borneo, C Kalimantan, Sintang, 100 m alt. Branches sparse puberulous. Leaves: blades elliptic or obovateelliptic, 13–16 by 3–4.5 cm, subcoriaceous, glabrous above, sparse puberulous on midrib and nerves beneath, apex cuspidate or caudate, base broadly cuneate; nerves 3–5 pairs, flat above, prominent beneath, nervules subparallel, conspicuous on both sides. Stipules triangular, acute, up to 4–5 mm long, puberulous. Cymes sessile, bracts absent. Flowers with 1 mm long pedicels, crowned; calyx campanulate, calyx mouth wide, truncate, with 5 indistinct teeth, pubescent. Drupes hemispheric, c. 2 mm diam, subglabrous or sparsely puberulous, 10-ribbed, crowned by the calyx limb; pyrenes 10. Habitat — Mixed dipterocarp forest. Distribution — Borneo, C Kalimantan. Note — The variety differs from the typical variety by having smaller leaves with nerves 3–5 pairs, 13–16 by 3–4.5 cm; sparse puberulous on branches, midrib and nerves beneath; smaller drupes c. 2 mm diam, with 1 mm long pedicels. 95. Lasianthus pergamaceus King & Gamble Lasianthus pergamaceus King & Gamble (1904) 123; Ridl. (1923a) 164; K.M. Wong (1989) 369, p.p. — Type: Wray 270 (holo SING; iso K), Peninsular Malaysia, Perak, 4500 ft. Lasianthus coriaceus King & Gamble (1904) 123. — Lectotype (designated by Wong 1989): Scortechini 332 (lecto SING; isolecto K), Perak; Ridley 10217 (para, n.v.), Peninsular Malaysia, Selangor. Lasianthus crassifolius Ridl. (1918) 85; (1923a) 165. — Lectotype (designated here): Ridley 15689 (lecto K), Peninsular Malaysia; Selangor; Maingay s.n. (para, n.v.), Peninsular Malaysia. Treelets; branchlets terete, robust, glabrous or minutely puberulous in the youngest part, sulcate when dry. Leaves: blades oblong-elliptic, 18–25 by 5–8 cm, thinly coriaceous, glabrous on both surfaces except minutely puberulous in young leaves beneath, apex cuspidate-acuminate, base broadly cuneate to subrounded, slightly asymmetric, midrib and nerves distinct or slightly depressed above, strongly prominent beneath, nerves 10–12 pairs, ascending at an angle of over 70°, curved to the margin, secondary nerves from the costa between nerves conspicuous, nervules distinct on both surfaces and prominent beneath, reticulate; petioles 7–10 mm long. Stipules triangular, 3–4 mm long, glabrous. Cymes shortly pedunculate; peduncles robust, 2–3 mm long, glabrous; bracts inconspicuous. Flowers sessile; calyx obconic-cupular, 3–5 mm long, with a truncate or minutely 4-toothed limb, glabrous; corolla tubular, pubescent outside, villous inside, 4-toothed; anthers 4. Drupes cupularglobose, c. 5–6 mm diam, glabrous or minutely hairy, crowned by a truncate or minutely 4-toothed limb; pyrenes 4. Distribution — Peninsular Malaysia, Philippines (Palawan). Note — This species is closely related to L. chinensis and L. repoeuensis, but it differs from both in having short robust peduncles and usually sulcate branchlets and cupulate calyx with truncate limb and drupe with 4 pyrenes. It is also similar to L. subaureus, but differs from the latter by having large leaves, sulcate branchlets, and peduncled cymes. 80 Blumea – Volume 57 / 1, 2012 Fig. 34 Lasianthus sabahensis H.Zhu. a. Habit with fruit; b. detail of node showing stipule and axillary inflorescences; c. calyx; d. corolla split opened; e. cross section of ovary; f. fruit; g. cross section of fruit (all: Chew & Corner RSNB 4598, holo L). 3 mm 2 cm 2 mm 3 mm 3 mm 81 H. Zhu et al.: Revision of Malesian Lasianthus Lasianthus humilis Elmer (1908) 519; Merr. (1923) 566. — Type: Elmer 9687 (holo lost; iso A, E, K, NY), Negros, Province of Negros Oriental, Cuernos Mountains, Dumaguete, Mar. 1908. Shrubs, 1–1.5 m high; branchlets terete, robust, 2–3 mm diam, compressed and enlarged at nodes, glabrous or subglabrous. Leaves: blades obovate-oblong, obovate or oblong, 10–18 by 3.5–7 cm, coriaceous, glabrous above, hirsute on midrib, nerves, and nervules beneath, apex shortly cuspidateacuminate, base oblique, obtuse, slightly cordate or rounded, midrib and nerves flat above, prominent beneath, nerves 8–9 pairs, ascending at an angle of 50–60°, curved to the margin; nervules distinct above, prominent beneath, parallel; petioles 3–7 mm long, sparsely hirsute. Stipules persistent, triangular or subulate, 5–6 mm long, hirsute. Cymes sessile; bracts linear, linear-lanceolate to lanceolate, 9–12 mm long, numerous, hirsute. Flowers fascicular, sessile; calyx campanulate, 2–3 mm long, sparsely hirsute outside, with 5 triangular 1 mm long teeth; corolla c. 10 mm long, puberulous outside, pubescent inside in the upper part. Drupes globose, glabrous, 4–5 mm diam, crowned by hirsute calyx teeth; pyrenes 5, smooth on the abaxial face. Distribution — NE India, China (S Yunnan), Indonesia (Sumatra, Java), Philippines (Negros Oriental). Note — This species has leaves obovate-oblong or obovate, 12–18 cm long, 5–7 cm broad, shortly cuspidate-acuminate at apex, the base oblique, obtuse, slightly cordate or rounded; stipules triangular or subulate, 5–6 mm long, hirsute; cymes sessile; bracts linear, linear-lanceolate to lanceolate, 9–12 mm long, numerous, hirsute; drupes globose, glabrous with 5 pyrenes. 107. Lasianthus robinsonii Ridl. Lasianthus robinsonii Ridl. (1909) 37; (1923a) 169; K.M.Wong (1989) 373. — Type: Ridley 13631 (holo SING; iso K), Peninsular Malaysia, Pahang, 4000–5000 ft. Shrubs, 1–2 m high; branchlets terete, c. 2 mm diam, densely tomentose. Leaves: blades oblong, 12–16 by 3.5–5 cm, membranaceous or thinly coriaceous, glabrous above, pubescent hairs on midrib and nerves beneath, apex cuspidate-acute, base subrounded to rounded, midrib and nerves flat to slightly depressed above, prominent beneath, nerves 8–9 pairs, ascending at an angle of 60° or more, curved to the margin, nervules distinct above, prominent beneath, subparallel; petioles 2–3 mm long, tomentose. Stipules triangular-lanceolate, 4–5 mm long, tomentose. Cymes long-pedunculate; peduncles slender, up to 5–6 cm long, tomentose; bracts 2, filiform, up to 8 mm long, tomentose; bracteoles like bracts but smaller, numerous. Flowers sessile; calyx c. 2.5 mm long, tomentose, lobes 4, linear, longer than tube; corolla c. 6 mm long, hairy both inside and outside, tube short, lobes oblong, longer than the tube. Drupes obovoid, tomentose to subglabrous, crowned by long linear calyx lobes; pyrenes 4. Distribution — Endemic to Peninsular Malaysia. 108. Lasianthus rotundatus Stapf Lasianthus rotundatus Stapf (1894) 185; Merr. (1921) 578. — Type: Haviland 1141 (holo K), Borneo, 8300 ft. Shrubs, c. 1 m high; branchlets terete, c. 1.5 mm diam, glabrous, slightly compressed in the youngest part, darkish when dry. Leaves: blades oblong-rounded, subrounded or rounded, 3–6 by 2–4.5 cm, coriaceous, glabrous on both surfaces, apex shortly cuspidate or apiculate, base rounded, blackish when dry, midrib and nerves prominent on both surfaces or slightly depressed above, nerves c. 5 pairs, ascending at an angle of 70–80°, curved to the margin, nervules prominent and distinct on both surfaces, subparallel; petioles 3–6 mm long, glabrous. Stipules triangular, c. 1 mm long, glabrous or sparsely puberulous. Cymes sessile or subsessile (peduncles less than 2 mm long); bracts absent. Flowers sessile; calyx campanulate, c. 2 mm long, tube c. 1 mm long, glabrous, limb c. 1 mm long, puberulous outside, teeth 4, broadly triangular, c. 0.6 mm long; corolla tube c. 4 mm long, lobes 4, ovate, c. 2.5 mm long; stamens 4; ovary 4-locular. Drupes subglobose, 3–4 mm diam, glabrous; pyrenes 4, smooth on the abaxial face. Distribution — Endemic to Borneo (Sabah). 109. Lasianthus sabahensis H.Zhu, sp. nov. — Fig. 34 Similis L. sterophyllo Merr., sed limbis calycis 4 minute dentibus, ovariis 3-locularibus, pyrenis 2 raro 3 differt. — Typus: Chew & Corner RSNB 4598 (holo L; iso K), Malaysia (Sabah), Mt Kinabalu, 5000 ft, Bembangan River, 1.3.1964. Shrubs, up to 3 m high; branchlets terete, c. 2 mm diam, puberulous to subglabrous, more or less compressed in the youngest part. Leaves: blades oblong or lanceolate-oblong, 9–15 by 2–5 cm, coriaceous, glabrous above, puberulous hairs on midrib and nerves beneath, apex acute to acuminate or cuspidate, margin reflexed, base cuneate, midrib prominent on both surfaces, nerves depressed or flat above, prominent beneath, 7–8 pairs, ascending at an angle of 60–80°, curved to the margin, nervules loosely anastomosing, subparallel, slightly prominent on both surfaces; petioles 1–1.5 cm long, puberulous. Stipules triangular, 4–6 mm long, with an acuminate to cuspidate apex, puberulous. Cymes sessile or shortly tuberculate; bracts absent. Flowers sessile, fascicular; calyx campanulate, tube c. 1 mm long, limb c. 1–1.5 mm long, more or less expanded, with 4 minute teeth, puberulous on both sides; corolla tubular, c. 5 mm long, puberulous outside, densely hairy in the upper half of the tube and the lobes inside, lobes 4; ovary 3-locular. Drupes ellipsoid, 8–9 mm long, c. 5 mm diam, puberulous or glabrous, crowned by a cupular calyx limb with 4 minute teeth, pyrenes 2, rarely 3, with thick, hard wall, smooth on the abaxial face, between pyrenes cork-like structure. Distribution — Endemic to Borneo. Habitat & Ecology — In forest between 800–1800 m altitude. Flowering from September to January. Notes — This species is similar to L. sterrophyllus, but differs in having relatively large stipules (4–6 mm long), calyx limbs with minute teeth, 3-locular ovary and 2-, rarely 3-pyrened drupes with cork-like structure between pyrenes, while L. sterrophyllus has stipules which are 3–4 mm long, calyx with 4 triangular lobes which are 2 mm long, and drupes with 5 pyrenes. This species seems somewhat between Lasianthus and Saldinia morphologically. Saldinia has a 2-locular ovary which develops to a 1-pyrened drupe with a hard wall. Lasianthus has an ovary with 3 or more locules, which develops to a drupe with 1 or more pyrenes with a hard and thick wall. Lasianthus sabahensis has a 3-locular ovary which develops to a drupe with 2 rarely 3 pyrenes of hard and thick walls. 110. Lasianthus scalariformis King & Gamble Lasianthus scalariformis King & Gamble (1904) 131; Ridl. (1923a) 167; M.R. Hend. (1933) 107; K.M.Wong (1989) 369. — Type: Scortechini 411 (lecto K, designated here), Peninsular Malaysia, Perak; Wray 332 (para, n.v.), Peninsular Malaysia, Perak, 6700 ft. Shrubs or treelets, up to 4 m high; branchlets terete, c. 2 mm diam, thinly pubescent, dark in colour when dry. Leaves: blades lanceolate-oblong to elliptic-oblong, 10–15 by 2.5–3.5 cm, coriaceous, glabrous on both surfaces except sparsely pubescent hairs on midrib and nerves beneath, apex acuminate to acute, 82 Blumea – Volume 57 / 1, 2012 Fig. 35 Lasianthus setulosus H.Zhu. a. Habit; b. detail of node showing axillary inflorescences and lower surface of leaf; c. flower; d. corolla split opened; e. calyx; f. fruit; g. cross section of fruit (a–e: R. George et al. S.42843, holo L; f, g: W.M.A. Brooke 9115, L). 2 cm 3 mm 3 mm 3 mm 3 mm 83 H. Zhu et al.: Revision of Malesian Lasianthus base acute, midrib and nerves flat to slightly prominent above, strongly prominent beneath, nerves 7–8 pairs, ascending at an angle of 50–60°, curved to the margin, nervules prominent on both surfaces, densely anastomosing, parallel; petioles 5–10 mm long, pubescent. Stipules triangular, 2–3 mm long, pubescent. Cymes pedunculate; main peduncles 5–10 mm long, robust, pubescent or glabrous with 2 branches; branches 2–4 mm long, pubescent or glabrous; bracts minute, subulate, c. 2 mm long, pubescent; bracteoles c. 1 mm long. Flowers sessile; calyx tube obconic, c. 2–3 mm long, pubescent, lobes 5, oblong or ovate-oblong, 3–4 by 1.5 mm, sparsely hirsute outside, puberulous inside; corolla tubular, 5–6 mm long, tomentose outside, lobes 5, triangular-ovate; stamens 5; filaments very short c. 1.2 mm long. Drupes obovoid, 3.5–4 mm diam, glabrous, 5-ridged, crowned by persistent calyx lobes; pyrenes 5. Distribution — Peninsular Malaysia, Indonesia (Sumatra). 111. Lasianthus setulosus H.Zhu, sp. nov. — Fig. 35 Similis L. borneensis Merr., sed ramulis et foliis dense longiore hispidis, calycibus minoribus, lobis calycis ovato-triangulis, 1.5 mm longis, apice acutis differt. A L. piloso Wight foliis basi subrotundatis, bracteis inconspicuis, lobis calycis ovato-triangulis, 1.5 mm longis differt. — Typus: R. George et al. S.42843 (holo L; iso K), Borneo, Limbang, on lower slope, 11 Sept. 1980. Shrub or treelet, 2–2.5 m high; branchlets terete, slightly compressed, c. 2 mm diam, densely dark-brown hispid. Leaves: blades elliptic-lanceolate, 10–17 by 2.5–5 cm, chartaceous to subcoriaceous, densely dark-brown hispid both surfaces, apex acuminate, base subrounded, slightly asymmetric, midrib depressed above, prominent beneath, densely hispid both surfaces or lower surface in old leaves, nerves slightly depressed and obscure above, prominent beneath, 9–12 pairs, ascending at an angle of c. 70°, curved to the margin, hispid, nervules obscure above, conspicuous beneath, subreticulate, hispid; petioles 0.5–1 cm long, densely hispid. Stipules subulatetriangular, c. 2 mm long, hispid. Inflorescences sessile, bracts inconspicuous. Flowers sessile, 5-merous; calyx campanulate, 3 by 3 mm, hispid, lobes 5, ovate-triangular, 1.5 by 1.5 mm, sparsely hispid both surfaces; corolla 6 mm long, tube puberulous external surface, lobes 5–6, lanceolate, 2 mm long, hirsute external surface, floccose hairs on internal surface, densely white lanate at the mouth. Drupes globose, 4–5 mm diam, hispid, 5-ridged, crowned by the persistent 5 ovate-triangular calyx lobes; pyrenes 5. Distribution — Borneo (Sarawak, Sabah, Brunei). Habitat & Ecology — In forest. Flowering from September to November. Additional specimens examined. Sarawak, G. Mulu, P.Chai S.39676 (L); Sg. Entulu, B. Lee S.54731 (L); Belarga, W.M.A. Brooke 9115 (L). Brunei, Temburong, C. Hansen 1622 (L), I. Nielsen & H. Balslev 1086 (AAU); Sukang, D.W. Kirkup & S. Thomas 619 (AAU). Sabah, Apensiangan Kayu, A. Mantor SAN 136864 (L). Note — This species is similar to L. borneensis. It differs from the latter by having densely spreading long hispid on branchlets and leaves on both surface, calyx relatively small, 3 by 3 mm, calyx lobes ovate-triangular with acute apex; while L. borneensis has very short hispid (strigose) on branchlets and leaves beneath, calyx 5 by 5 mm, calyx lobes oblong with obtuse apex. It differs from L. pilosus by having leaves with subrounded base, bracts inconspicuous, calyx lobes ovatetriangular, 1.5 mm long. Lasianthus pilosus has leaves with subcordate base; bracts linear, 5 mm long; calyx lobes linear, up to 5 mm long or more. 112. Lasianthus sikkimensis Hook.f. Lasianthus sikkimensis Hook.f. (1880) 180; Deb & M.Gangop. (1991) 297; H.Zhu (1994) 60; (1998) 155; H.S.Lo (1999) 80, pl. 21: 6–11. — Type: Hooker & Thomson 11 (holo K), Sikkim-Himalaya. Lasianthus mindanaensis Merr. (1922) 466; (1923) 567. — Type: Bur. Sci. 38795 (Ramos & Endano s.n.) (holo lost; iso K, L, US), Philippines, Mindanao, Mt Candoon, syn. nov. Lasianthus langkokensis (Drake) Pit. (1924) 379. — Mephitidia langkokensis Drake (1895) 240. — Lasianthus sikkimensis Hook.f. subsp. langkokensis (Pit.) H.Zhu (1994) 61; (1998) 153. — Type: Balansa 2654 (holo P), Vietnam, Tonkin, Bavi. Lasianthus baviensis (Drake) Pit. (1924) 380. — Mephitidia baviensis Drake (1895) 239. — Type: Balansa 2655 (holo P; iso K), Vietnam, Tonkin, Tu-phap. Lasianthus tsangii Merr. ex Li (1943) 458; H.J.Li & H.Y.Liu (1998) 295. — Type: Tsang 23940 (holo SCBI), China, Kwangsi [Guangxi]. Shrubs, 1–3 m high; branches and branchlets densely browntomentose. Leaves: blades elliptic-lanceolate or elliptic, 10–20 by 2.5–5 cm, subcoriaceous or thinly coriaceous, glabrous above, densely brown-tomentose beneath, apex cuspidateacuminate, base acute or obtuse, usually recurved at margin, midrib and nerves flat above, conspicuously prominent beneath, nerves 7–10 pairs, ascending at an angle of over 50°, curved to the margin, nervules subparallel, prominent beneath; petioles 6–14 mm long, brown-tomentose. Stipules triangular, 2–4 mm long, densely tomentose. Cymes sessile, 1–3-flowered; bracts persistent, linear to lanceolate, up to 2 cm long, brown-tomentose. Flowers sessile or subsessile; calyx 3–4 mm long, tube glabrous outside, lobes 5, triangular, hirsute outside; corolla c. 10 mm long, puberulous outside. Fruits ellipsoid, 6–8 mm long, glabrous except on the persisting hirsute calyx lobes; pyrenes 5. Distribution — NE India, Sikkim, Bangladesh, China, Vietnam, Thailand, Philippines. Note — Lasianthus mindanaensis has branches rather densely brown-tomentose; leaves chartaceous to subcoriaceous, lanceolate, 11–18 cm long, apex caudate, base acute, nerves 7, nervules rather distinct beneath, yellowish brown-tomentose on nerves and nervules beneath, petioles densely brown-tomentose; stipules ovate, 3–4 mm long, densely brown-tomentose; flowers sessile, fascicles, bracts narrowly lanceolate, 8 mm long, brown-tomentose; calyx tube glabrous, lobe 2 mm long, glabrous outside; drupes glabrous, 5 mm long. All of these characters agree well with L. sikkimensis. Lasianthus mindanaensis is conspecific with L. sikkimensis. 113. Lasianthus sogerensis Wernham Lasianthus sogerensis Wernham (1918) 72. — Type: Forbes 313 (holo BM), New Guinea, Sogere, 3000 ft. Shrubs, up to 2 m high; branchlets terete, c. 2 mm diam, densely spreading villous. Leaves: blades oblong to obovate-oblong, 7–11 by 2–4 cm, chartaceous, sparsely villous above but densely villous hairs on midrib, densely villous beneath, apex cuspidate-acuminate, base subrounded to rounded, more or less oblique, the margin ciliate, nerves obscure above, prominent and distinct beneath, 8–11 pairs, ascending at an angle of 50–60°, curved to the margin, nervules obscure above, distinct beneath, subparallel; petioles 2–5 mm long, densely villous. Stipules inconspicuous. Cymes sessile, bracts paired in each leaf axil, leaf-like, ovate-lanceolate to lanceolate, up to 1.5 cm long, nerved, long-villous, apex caudate with a linear tip. Flowers sessile; calyx tube campanulate, c. 3 mm long, densely villous, lobes 4, linear-lanceolate, c. 3 mm long, villous; corolla tubular, c. 5 mm long, villous outside. Drupes globose, villous, c. 4 mm diam, 4–5-ridged, verrucose, crowned by linear-lanceolate calyx lobes; pyrenes 4–5, verrucose on the abaxial face. Distribution — Indonesia (Moluccan, Irian Jaya) and Papua New Guinea. Note — This species has densely spreading villous hairs on branches, leaves, stipules, bracts, and drupes. It has chartaceous leaves with subrounded, or rounded and slightly oblique, 84 Blumea – Volume 57 / 1, 2012 Fig. 36 Lasianthus solomonensis H.Zhu. a. Habit; b. detail of lower surface of leaf showing indumentum; c. detail of node showing stipule and axillary inflorescences; d. flower; e. corolla split opened; f. fruit; g. cross section of fruit (all: T.C. Whitmore RSS6051, holo L). 2 cm 2 mm 2 mm 2 mm 85 H. Zhu et al.: Revision of Malesian Lasianthus bases, and ciliate margin, 8–11 pairs of nerves. Its stipules are inconspicuous but bracts are paired, leaf-like, ovate-lanceolate, with a long linear apex. It has drupes with 4–5 pyrenes crown by linear-lanceolate calyx lobes. This species is closely related to L. attenuatus and L. cyanocarpus, but differs from both of them by having densely patent or spreading villose hairs on the plant, bracts with a long linear apex. Furthermore, it differs from L. attenuatus by leaf-like paired bracts, and differs from L. cyanocarpus by ovate-lanceolate bracts paired in each leaf axil. Wernham (1918) did not mention bracts but described stipules with two long prongs (8 mm long), and also described the drupes with 2 pyrenes. In our observation of the specimens, which match type well, this species has inconspicuous stipules but conspicuously ovate-lanceolate bract pairs, and drupes with 4 pyrenes. Wernham could have called the bracts stipules. 114. Lasianthus solomonensis H.Zhu, sp. nov. — Fig. 36 Proximus L. chlorocarpus K.Schum., sed ramulis dense pubescentibus, floribus 5–7 meris, lobis calycis triangulato-subulatis aequalis tubo, drupis minoribus, pyrenis 5–7 differt. A L. clementis Merr. foliis majoribus, stipulis lineari-subulatis 5 m longis, floribus 5–7-meris, bracteis subulatis 2 mm longis, lobis calycis aequalis tubo differt. — Typus: Whitmore RSS6051 (holo L), Solomon Islands, Guadalcanal, Mt Gallego, 2000 ft alt., in very steep unstable hillside, 6 July 1965. Shrub, c. 1–2 m high; branchlets terete or compressed in the youngest part, c. 1.5 mm diam, densely appressed brownish pubescent. Leaves: blades elliptic-oblong, 10–12 by 3–4 cm, subcoriaceous, glabrous above, puberulous beneath, appressed-pubescent midrib and nerves beneath, apex cuspidate, with c. 2 mm long arista, margin slightly undulate and recurved, base acute; midrib flat or slightly prominent above, prominent beneath, nerves slightly prominent above, prominent beneath, 6–7 pairs, ascending at an angle of 60–70°, curved to margin, nervules conspicuous beneath, parallel; petioles 0.8–1 cm long, densely pubescent. Stipules linear-subulate, c. 5 mm long, densely pubescent. Inflorescences sessile; bracts small, subulate, c. 2 mm long, pubescent. Flowers fascicular, sessile; calyx campanulate, c. 3 mm long, pubescent, lobes 5–7, triangular-subulate, c. 1.5 mm long, pubescent external surface, glabrous internal surface; corolla c. 5 mm long, pubescent external surface, lobes 5–7, lanceolate; stamens 5–8, filament very short; styles c. 4 mm long, with 5–7 linear stigmas. Drupes globose, c. 3 mm diam, pubescent, crowned by 5–7 subulate calyx lobes; pyrenes 5–7, slightly verrucose on abaxial face, ovate on the transverse section. Distributed — Solomon Islands. Habitat & Ecology — In primary forest between 300–2000 ft altitude. Flowering and fruiting between July and December. Note — This species is closely related to L. chlorocarpus, but differs from the latter by having branchlets densely pubescent, flowers 5–7-merous, calyx lobes triangular-subulate, as long as calyx tube, drupes relatively small with 5–7 pyrenes. Lasianthus chlorocarpus has branchlets and leaf nerves beneath strigose, flowers 4-merous, calyx usually with a limb truncate or small dentate, drupes relatively big, pyrenes 4. This species differs from L. clementis by having leaves relatively big, stipules linearsubulate, 5 mm long, flowers 5–7-merous, bracts subulate, 2 mm long, calyx lobes as long as calyx tube, drupes with 5–7 pyrenes. 115. Lasianthus stercorarius Blume Lasianthus stercorarius Blume (1826–1827) 1000; Miq. (1857) 324; (1869) 248; Merr. (1921) 577; Bakh.f. (1965) 341. — Type: Blume s.n. (holo L, barcode L0000707), Java, Mt Tjerimai. Lasianthus stercorarius Blume var. polypleura Blume ex Miq. (1869) 248, syn. nov. — Type: Blume s.n. (lecto L, designated here, barcode L0305599), Java; Blume s.n. (para L, barcode L0305583), Java. Lasianthus mucronulatus (Korth.) Miq. var. lineata Miq. (1869) 249, syn. nov. — Type: Korthals s.n. (holo L, barcode L0310824), Sumatra. Shrubs to treelets, up to 4.5 m high; branchlets terete, c. 2 mm diam, thinly appressed-puberulous or glabrous. Leaves: blades oblong-lanceolate to ovate-lanceolate, 7–15 by 2–4 cm, subcoriaceous to coriaceous, glabrous above, puberulous hairs on venation beneath or glabrous also, apex acuminate-caudate, base acute, midrib slightly depressed above, prominent beneath, nerves flat and slightly obscure above, prominent beneath, 5–9 pairs, ascending at an angle of 45–60°, curved to the margin, nervules obscure above, distinct beneath, subparallel; petioles 5–10 mm long, puberulous or glabrous. Stipules triangular, 3–6 mm long, coriaceous, swollen at the base, puberulous or glabrous. Cymes sessile; bracts absent. Flowers sessile; calyx urceolate or urceolate-cupular, c. 1.5–2 mm long, with a truncate or minutely toothed limb, puberulous or glabrous; corolla tubular, c. 5–7 mm long, puberulous outside. Drupes globose, glabrous or thinly puberulous, 3–5 mm diam, acute at the base, truncate or minutely toothed at the apex, conspicuously 6–9-ridged; pyrenes 6–9, smooth on the abaxial face. Distribution — Malaysia (Peninsular, Borneo) and Indonesia (Sumatra, Java, Kalimantan, Sulawesi). Habitat & Ecology — In primary forest. Notes — Lasianthus stercorarius is a very variable species in leaves, indumentum, and drupes. It is keyed out mainly by its conspicuously ovate-triangular stipules with swollen bases. The type specimen shows thin puberulous hairs on branches and leave nerves beneath; subcoriaceous leaves with 5–6 nerves and nervules reticulate, apex acuminate, base acute; calyx limbs subtruncate and with 4–5 minute teeth; drupes with 8–9 pyrenes. Two varieties were established: var. polypleura based on specimens with leaves of 8 pairs of nerves; var. palembanicus based on specimens with leaves of 12–14 pairs of nerves. The variety palembanicus is conspecific to L. mucronulatus and should be reduced to a synonym of the latter. Lasianthus stercorarius differs from L. mucronulatus by having usually relatively small subcoriaceous leaves with acute to cuneate base and less than 10 pairs of nerves and conspicuously more than 2 mm long triangular stipules with swollen bases. Lasianthus mucronulatus var. lineata was established on Korthals’s Sumatra specimens. It has triangular stipules with swollen base, subcoriaceous leaves with much acute base and should be reduced to L. stercorarius. 116. Lasianthus sterrophyllus Merr. Lasianthus sterrophyllus Merr. (1937) 299. — Type: Winkler 508 (holo HBG; iso NY), Borneo, Bukit Mulu, 950 m alt. Shrubs or treelets, up to 3 m high; branchlets terete, 2–3 mm diam, glabrous except sparsely hirsute at nodes. Leaves: blades oblong-lanceolate, 8–13 by 2.5–3.5 cm, coriaceous, glabrous above, glabrous or sparsely hirsute hairs on midrib and nerves beneath, apex acute to caudate-acute, base acute, midrib and nerves slightly prominent above, prominent beneath, nerves 6–9 pairs, ascending at an angle of 55–70°, curved to the margin, nervules prominent on both surfaces, subparallel; petioles 5–10 mm long, glabrous or sparsely hirsute. Stipules triangular, 3–4 mm long, coriaceous, glabrous or pubescent. Cymes sessile; bracts absent or inconspicuous. Flowers sessile calyx campanulate, subglabrous, tube c. 2 mm long, lobes 4, triangular, c. 2 mm long, sparsely hirsute; corolla not seen. Drupes subglobose, c. 4 mm diam, glabrous or sparsely hirsute, crowned by persistent triangular calyx lobes; pyrenes 5, smooth on the abaxial face. Distribution — Malaysia (Sarawak), Indonesia (Kaliman tan). 86 Blumea – Volume 57 / 1, 2012 Notes — This species is similar to L. inodorus subsp. montigenus. Lasianthus inodorus subsp. montigenus has conspicuous orbicular or ovate bracts (although they are usually caducous, you can always see some in most specimens) and is found above 1700 m altitude. Lasianthus sterrophyllus has inconspicuous bracts and has 4 ovate 2 mm long calyx lobes, and is usually found below 1700 m altitude. An imperfect specimen in Leiden (Winkler 508) differs from the type description of L. sterrophyllus in having conspicuous ovate bracts. The label on the specimen looks quite new from the paper appearance and typing writing, and the date of collection 2/12/1925 does not agree with Merrill’s type Winkler 508. The specimen obviously is L. inodorus subsp. montigenus. 117. Lasianthus stipularis Blume Lasianthus stipularis Blume (1826–1827) 997; Miq. (1857) 319; (1869) 246; Kurz (1877) 30; Hook.f. (1880) 179; King & Gamble (1904) 112; Elmer (1906b) 10; Ridl. (1923a) 152; Merr. (1929a) 292; Craib (1934) 217; K.M.Wong (1989) 368; H.Zhu (2001b) 141. — Mephitidia stipularis Korth. (1851) 223. — Type: Blume s.n. (holo L, barcode L0000708; iso L, barcode L0000709), Java, Salak. Lasianthus stipularis Blume var. novoguineensis Valeton (1911) 498, syn. nov. — Type: Versteeg 1228 (holo L; iso K), SW New Guinea. Shrubs, 1–2 m high, entirely glabrous except bracts and corolla; branchlets terete, 2–3 mm diam, compressed in the youngest part. Leaves: blades oblanceolate-oblong, 12–16 by 3–5.5 cm, membranaceous, greenish brown when dry, apex cuspidate-caudate, base cuspidate-cuneate, midrib and nerves flat to slightly prominent above, prominent beneath, nerves 9–10 pairs, slender, ascending at an angle of over 50°, curved to the margin, nervules faint, subparallel; petioles 5–10 mm long; stipules broadly triangular-ovate, ovate or ovate-orbicular, larger, covering the cymes, up to 1.25 cm long and broad at base, membranaceous, glabrous, persistent. Cymes sessile; bracts numerous, linear to narrowly lanceolate, villous, up to 1.5 cm long, often exceeding the stipules. Flowers sessile; calyx campanulate, teeth broadly triangular, c. 1 mm long, hirsute; corolla not seen. Drupes ovoid-globose, glabrous, 4–5 mm diam, crowned with calyx limbs, with 4 obtuse ridges; pyrenes 4. Distribution — Burma, Thailand, Malaysia (Peninsular, Borneo), Brunei, Indonesia (Sumatra, Java, Kalimantan, Sulawesi, Moluccas), Philippines, Papua New Guinea. Note — This species has numerous hairy linear to narrowly lanceolate bracts up to 1.5 cm long, in which it differs from L. pseudostipularis (bracts minute and inconspicuous). Lasianthus stipularis var. novoguineensis differs from the typical variety by only having oblong to elliptic-lanceolate leaves, which is not enough to separate it from L. stipularis as a variety of the latter. 118. Lasianthus subaureus Craib Lasianthus subaureus Craib (1933) 27; (1934) 218; K.M.Wong (1989) 368; H.Zhu (2001b) 142. — Type: Kerr 14117 (holo K; iso BK), Thailand, Puket. Shrubs, up to 3 m high; branchlets terete, c. 2 mm diam, densely appressed puberulous, aureate when young. Leaves: blades oblong-lanceolate, 10–17 by 1.7–3.5 cm, coriaceous, glabrous, luteous above, pubescent hairs on midrib and nerves, or nervules beneath, apex acuminate, margin often reflexed, base acute to cuneate or rounded, slightly unequal-lateral, midrib conspicuous, more or less depressed above, prominent beneath, nerves prominent on both surfaces, c. 12 pairs, ascending at an angle of over 50°, curved to the margin, nervules distinct and slightly prominent on both surfaces, reticulate; petioles 10–15 mm long, densely appressed puberulous. Stipules triangular-subulate, 2.5–3 mm long, puberulous. Cymes sessile or rarely extremely congested; bracts absent. Flowers sessile; calyx cupulate, c. 4 mm long, appressed puberulous, tube 1.5–2 mm long, limb cupular, apex truncate or minutely teethed; corolla tubular, pubescent outside, tube up to 12 mm long, lobes 6, linear or linear-lanceolate, pilose outside; ovary with 6 locules. Drupes subglobose, 4–5 mm diam, puberulous, conspicuously 5–6-ridged, base narrowed, apex crowned by a cupular and truncate or minutely toothed calyx limb; pyrenes 5–6, smooth on the abaxial face. Distribution — Burma, Thailand, Peninsular Malaysia. Note — This species is close to L. verticillatus by its cupular calyx with truncate limb. It differs from the latter by having relatively narrow leaves with reticulate veinlets and longer petioles. 119. Lasianthus subglobosus H.Zhu, sp. nov. Similis L. bruneensis H.Zhu, sed ramulis dense depressi-pubescentibus, foliis basi acutis, nervulis conspicuis, petiolis pubescentibus 5 mm longis et pyrenis 6 differt. — Typus: Puasa-Angian 3967 (holo K; iso L), Malaysia (Sabah), Tambunan, 1200 ft, 19/02/1934. Treelet, 2 m high; branchlets terete, densely depressed-pubescent. Leaves: blades lanceolate to oblong-lanceolate, 8–12 by 1.8–4.5 cm, coriaceous, pubescent on leaf nerves beneath, apex acuminate or caudate, base acute, midrib and nerves conspicuously prominent on both surfaces, nerves 5–7 pairs, ascending at an angle of 50–60°, curved to the margin, nervules conspicuous on both surfaces, reticulate; petioles 0.5 cm long, pubescent. Stipules small, triangular, 1.5–3 mm long, pubescent. Inflorescences sessile, bracts inconspicuous. Flowers fascicular, sessile; calyx campanulate, 2 mm long, pubescent, calyx lobes 5, triangular, 1–2 mm long; corolla not seen. Drupes c. 3 mm diam, subglobose, puberulous, black, crowned by persistent small subulate-triangular calyx teeth; pyrenes 6, shallow sulcate on the abaxial face, triangular in transverse section. Distribution — Endemic to Borneo. Habitat & Ecology — In forest. Flowering and fruiting from February to December. Note — This species is close to L. bruneensis, but differs from the latter in having branchlets densely depressedpubescent; leaves with an acute base, nervules conspicuous on both surfaces; petioles 5 mm long, pubescent; drupes with 6 pyrenes. 120. Lasianthus submembranifolius Elmer Lasianthus submembranifolius Elmer (1913) 1871; Merr. (1923) 567. — Type: Elmer 14261 (iso A, CAS, K, L, MO, NY), Philippines, Mindanao, Mt Urdaneta, 3250 ft. Shrubs; branchlets terete, c. 2 mm diam, compressed in the youngest part, glabrous. Leaves: blades ovate-oblong, 10–15 by 3–5 cm, more or less unequal-lateral, thinly membranaceous, glabrous above, minutely puberulous hairs on midrib and nerves beneath, apex cuspidate-acute, base cuneate, asymmetric, midrib and nerves almost flat above, prominent beneath, nerves 4–5 pairs, ascending at an angle of 40–50°, curved toward the margin and joining the nerves above very near the margin, forming an obscure intramarginal vein, nervules distinct beneath, loosely anastomosing, reticulate; petioles less than 1 cm long, glabrous. Stipules broadly ovate, c. 2 by 3 mm, glabrous. Cymes tuberculate; bracts minute, numerous, subulate, 0.5–1 mm long, glabrous. Flowers shortly pediceled; pedicels 1–2 mm long, glabrous; calyx shallow-cupular, 1–2 mm long, glabrous, with 5 oblong-triangular lobes; corolla c. 1.5–3 mm long, glabrous outside, with 5 bluntly oblong lobes; stamens 5; 87 H. Zhu et al.: Revision of Malesian Lasianthus Fig. 37 Lasianthus sumatraensis H.Zhu. a. Habit with inflorescences; b. detail of inflorescence; c. calyx; d. fruit; e. cross section of fruit (all: W.J.O. de Wilde et al. 13225, holo L). 2 cm 2 mm 5 mm 5 mm 88 Blumea – Volume 57 / 1, 2012 filament short. Drupes subglobose, up to 7.5 mm long; pyrenes 5, coarsely rugose on the abaxial face. Distribution — Endemic to the Philippines. Note — According to Elmer’s description, this species has rugose drupes with 5 pyrenes. 121. Lasianthus sumatraensis H.Zhu, sp. nov. — Fig. 37 Similis L. malaiense H.Zhu, sed foliis basi acutis praeter costae et nervis subtus glabris, cyma ramosa, floribus pedicellatis, lobis calycis linearis 5 mm longis differt. A L. scalariformis King & Gamble ramulis dense brunneotomentosis, costis et nervis subtus brunneo-tomentosis, foliis ovato-ellipticis et lobis calycis linearibus tomentosis differt. — Typus: De Wilde et al. 13225 (holo L), North Sumatra, Kutatjane, 2500 m alt., in montane mossy forest, 23 June 1972. Shrub, up to 4 m high; branchlets terete, densely brown tomentose. Leaves: blades ovate-elliptic, 7–11 by 2.5–4.5 cm, coriaceous, glabrous except venation beneath, apex acute to acuminate, margin ciliate, base acute to subrounded, midrib depressed above, prominent beneath, brown tomentose beneath, nerves flat above, prominent beneath, 6–7 pairs, ascending at an angle of 50–60°, curved to the margin, tomentose beneath, nervules prominent on both surfaces, conspicuous, parallel, pubescent to glabrescent beneath; petioles 1–1.5 cm long, brown tomentose. Stipules lanceolate, 3–4 mm long, tomentose. Inflorescences cymose, brown tomentose; the main peduncles 5–10 mm long, peduncles branches (if present) c. 5 mm long; bracts linear, 2 in each node, 5 mm long, brown hirsute; bracteoles 2, linear, 3–5 mm long, brown hirsute. Flowers pedicellate; pedicels 1–3 mm long, brown tomentose; calyx campanulate, densely brown tomentose, lobes 5, linear, 5–6 mm long, densely tomentose external surface. Drupes subglobose, 5-ridged, crowned by persistent linear, 5 mm long, calyx lobes; pyrenes 5, smooth on the abaxial face, suborbicular in transverse section. Distribution — Sumatra (northern part). Habitat & Ecology — In montane rainforest to mossy forest between 1600–2600 m altitude. Flowering and fruiting between May and August. Additional specimens examined. North Sumatra, Atjeh, 2400–2600 m alt., W.J.O. de Wilde et al. 16785 (K, L); Atjeh, 1600–1800 m alt., K. Iwatsuki et al. S.958; Atjeh, Van Steenis 9545 (L). Note — This species is quite similar to L. malaiense. It differs from the latter by having leaves with acute base, glabrous except venation beneath; cymes branched; flowers pedicellate; calyx lobes linear, 5 mm long. While L. malaiense has leaves with rounded base, villose beneath; cyme capitate; peduncles 2–3 cm long; flowers sessile; calyx lobes lanceolate, 2.5 mm long. It differs from L. scalariformis by having dense brown tomentose hairs on branchlets, leaf midrib and nerves beneath; ovate-elliptic leaves; calyx lobes linear, densely tomentose. While L. scalariformis has leaves subglabrous, elliptic-lanceolate; calyx lobes oblong-lanceolate, sparsely hirsute. 122. Lasianthus sylvestroides Valeton Lasianthus sylvestroides Valeton (1927) 109; Merr. & L.M.Perry (1946) 219. — Type: Ledermann 12655a (holo B†), New Guinea, Felsspitze, 1400– 1500 m; Brass 13025 (neo L, designated here), New Guinea, Idenburg River. Shrubs; branchlets terete, tomentose. Leaves: blades oblong or elliptic-lanceolate, 7–11 by 2.5–4 cm, subcoriaceous, glabrous above except tomentose midrib, tomentose beneath, apex acute to acuminate, mucronate, base acute, margin recurved, midrib flat above, prominent beneath, nerves flat to slightly depressed above, prominent beneath, 7–10 pairs, ascending at an angle of over 45°, curved to the margin, nervules prominent on both surfaces, parallel; petioles, c. 3–5 mm long, tomentose. Stipules linear-subulate, 4–5 mm long, tomentose. Cymes axillary, sessile or subsessile; bracts linear, 5–6 mm long, singular, tomentose. Flowers fascicular, sessile; calyx obconical, tube c. 2 mm long, hirsute, lobes 5, linear, 3–4 mm long, hirsute; corolla tomentose. Drupes globose, 4–5 mm diam, with 5–6 obtuse ridges, verrucose, crowned by the persistent linear to lanceolate-subulate, 3–4 mm long, hirsute calyx lobes; pyrenes 5–6. Distribution — Indonesia (Sulawesi, Irian Jaya), Papua New Guinea. Note — The specimen Brass 13025 so closely fits the description of Valeton’s species according to Merrill (Merrill & Perry 1946) that we select it as a neotype. 123. Lasianthus tenuifolius H.Zhu, sp. nov. — Fig. 38 Similis L. attenuato var. minore H.Zhu, sed foliis tenui-chartaceis, stipulis subulato-linearibus, bracteis inconspicuis, fructibus glabribus et pyrenis 7–8 differt. — Typus: W. Ave 100 (holo L), Peninsular Malaysia, Perak, 550 m alt., in hill dipterocarp forest, 31 Aug. 1982. Shrub, 1 m high; branchlets slender, terete, spreading villous. Leaves: blades narrow-oblong to linear-oblong, 6–10 by 1.5–2.5 cm, thinly chartaceous, sparsely spreading villous on both surfaces, densely on midrib and nerves, apex acute, margin ciliate, base rounded, oblique, midrib flat above, prominent beneath, nerves obscure above, slightly prominent beneath, 6–9 pairs, ascending at an angle of 45–50°, curved to the margin; petioles short, 1–2 mm long, villous. Stipules subulate-linear, 3–5 mm long, villous. Inflorescences sessile; bracts inconspicuous. Flowers fascicular; calyx campanulate, 3–4 mm long, glabrous to sparsely villous on external surface, lobes 5, triangular-lanceolate, 2.5 mm long, glabrous to sparsely villous; corolla 6–7 mm long, villous on external surface, lobes ovate-triangular. Drupes oblate-globose, 4–5 by 3–4 mm, glabrous, yellowish, 7–8 conspicuously sulcate; pyrenes 7–8, ovate-triangular in transverse section, smooth but with a furrow on the abaxial face, shallow furrow on the sides. Distribution — Malaysia (Peninsular, Borneo) and Indonesia (Sumatra). Habitat & Ecology — In hill rainforest between 50–600 m altitude. Flowering and fruiting between August and November. Additional specimens examined. W Kalimantan, Kohyama et al. K 10591 (L). Sarawak, P.Chai et al. S.31536 (L); C. Hose 156 (L). Brunei, M.J.E. Coode 6402 (L). Sabah, B.C. Stone SAN 85255 (K, L). Borneo, Bukit Raya, Veldkamp 8048 (L); Korth. s.n. W Sumatra, H. Nagamasu 3277 (L). Peninsular Malaysia, Selangor, KEP-FRI 1958 (L); Perak, W. Ave 24 (L); Pasoh, KEP-FRI 25703 (L); Pasoh, E.G.1532 (L). Note — This species is quite similar to L. attenuatus var. minor. It differs from the latter by having thinly chartaceous leaves; nerves obscure upper surface; stipules subulate-linear; bracts inconspicuous; drupes glabrous; pyrenes 7–8. Lasianthus attenuatus var. minor has chartaceous to subcoriaceous leaves with conspicuous nerves upper surface; stipules linear-lanceolate; bracts conspicuous, linear to linear-lanceolate; drupes hirsute; pyrenes 4–5. This species was confused with L. cyanocarpus (= L. inaequalis). Miquel (1857) doubted that a drupe with 8 pyrenes was found in L. inaequalis and later stated that drupes with 8 pyrenes do not occur in L. inaequalis (Miquel 1869). In our opinion, the specimens with 8-pyrened drupes present the new species. 124. Lasianthus tomentosus Blume Lasianthus tomentosus Blume (1826–1827) 997; Miq. (1857) 318, p.p. quoad type specimens; (1869) 245; Bakh.f. (1965) 343. — Type: Blume s.n. (lecto L, designated here, barcode L0057521), Java. Shrubs, c. 2 m high; branchlets terete, c. 2 mm diam, densely to thinly brown-tomentose. Leaves: blades ovate-oblong to 89 H. Zhu et al.: Revision of Malesian Lasianthus Fig. 38 Lasianthus tenuifolius H.Zhu. a. Habit; b. leaf; c. stipule; d. fruit; e. cross section of fruit (all: W. Ave 100, holo L). 1.5 cm 2 cm 5 mm 5 mm 5 mm 96 Blumea – Volume 57 / 1, 2012 5 mm long peduncles, bracts lanceolate, 4–7 mm long; Fruits hairy, with numerous small seeds. It is clearly a Urophyllum. Litosanthes brauniana Warb. ex Boerl. (1891) 142. — Lasianthus braunianus (Warb. ex Boerl.) M.Gangop. & Chakrab. (1992) 338. = Amaracarpus braunianus (Warb. ex Boerl.) Valeton (1911) 501. Acknowledgements This research was supported by a grant from NWO of the Netherlands (grant no. B 85-340) and grants of the National Natural Science Foundation of China (No. 30770158, 31170195, 41071040). We thank the Herbaria AAU, BK, BKF, BM, C, E, HBG, K, KEP, L, MO, NY, P, SING, U, US for checking specimens and loan of type specimens. We are grateful to Prof. P. Baas, Dr. F. Adema, Prof. P.C. van Welzen and Dr. W.J.J.O. de Wilde (all L) for their great help in the research, and to Dr. J.F. Veldkamp (L) for his great help in botanical nomenclature. We thank Mr. Wu Xilin, a botanical artist of the herbarium of Kunming Institute of Botany, the Chinese Academy of Sciences for illustrations in the article. We also thank reviewers and editors very much for their important and constructive comments. REFERENCES Andersson L, Rova JHE. 1999. The rps16 intron and the phylogeny of the Rubioideae (Rubiaceae). Plant Systematics and Evolution 214: 161–186. Baillon H. 1880. Histoire des plantes 7: 546. Hachette, Paris. Bakhuizen van den Brink Jr RC. 1963. Notes on the flora of Java VIII. 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Yunnan People’s Press, Kunming. 98 Blumea – Volume 57 / 1, 2012 Xiao LQ, Zhu H. 2007. Paraphyly and phylogenetic relationship in Lasianthus (Rubiaceae) inferred from chloroplast rps16 data. Botanical Studies 48: 227–232. Yamazaki T. 1964. Note on Lasianthus in Ryukyu. Journal of Japanese Botany 39: 348–352. Zhu H. 1994. A taxonomic study on the genus Lasianthus Jack in China. Acta Phytotaxonomica Sinica 32: 49–81. Zhu H. 1998. Notes on the genus Lasianthus Jack (Rubiaceae) from Asia. Acta Botanica Yunnanica 20: 149–159. 8053: 64 – BW series 4851: 21; 5549: 56; 7950: 56; 8406: 21; 10659: 113; 10675: 26; 12622: 49a; 13339: 73; 13531: 59; 13870: 26; 14278: 73 – Bygrave 35: 46. Carr 12000: 21; 12516: 26; 12517: 26; 12539: 21; 12648: 21; 12891: 21; 14365: 26; 14396: 121 – Carrick et al. 21: 53b; 1456: 117 – Chakrabarty 1220: 56; 2076: 56 – Chand 2769: 128; 4746: 49a; 4889: 60a; 5436: 60a; 5438: 60a; 6321: 56 – Chao 883: 49a; 884: 35; 893: 35 – Chew 64: 97a; 232: 82a; 362: 12; 764: 75; 790: 82a; 834: 3; 881: 72a; 1103: 121 – Chew et al. RSNB 974: 11; 1019: 27; 1033: 63; 1435: 63; 1718: 63 – Ching S55287: 53a; S55288: 14 – Chua FRI39084: 44 – Chuang & Kao 3280: 49a; 3439: 49a – Church et al. 10: 128; 198: 53a; 423: 94; 713: 94b; 870: 37; 888: 7; 937: 7; 1441: 68; 1451: 102; 1681: 71; 1789: 128; 1793: 53b; 1930: 53b; 2001: 67; 2051: 53a; 2245: 53a; 2403: 53a; 2424: 72b; 2639: 65; 2734: 37 – Clemens J. & Clemens M.S. 343: 21; 1192: 6a; 9467: 12; 26106: 53a; 26221: 63; 26271b: 53a; 26452: 63; 26463: 94; 26657: 78; 27353: 29; 27713: 60c; 27893: 78; 27893b: 60c; 27894: 130; 28105: 130; 28870: 130; 28876: 63; 29561: 130; 29578: 63; 29597: 109; 29691: 108; 29838: 92; 30348: 103; 30400: 53a; 31016: 103; 31479: 103; 32536: 60b; 32538: 42; 32916: 108; 34307: 42; 35158: 42; 40617: 11 – Coert 999: 26; 1423: 48 – Colfs 194: 48 – Collenette 21578: 42 – Collins 819: 128 – Conn 1719: 6a – Coode 5493: 117; 5846: 125a; 6402: 123; 6896: 72b – Craven & Schodde 1082: 49a; 1403: 131 – Curtis 284: 35; 2081: 87. Dakkus 264: 100 – Danser 5725: 106; 5743: 115; 5761: 101; 5773: 106; 5788: 64 – Dausus G.48: 128 – Davidse 7907: 103 – Davis 481: 17; 706: 56; 707: 59 – De Jong 760: 71; 889: 67 – De Silva s.n. ex Wall. Cat. 8442: 6a – De Vogel 886: 29; 1319: 94; 1739: 102; 1772: 117; 2766: 54; 2832: 101; 5124: 26; 5136: 92; 5344: 92; 8262: 103; 8266: 53a; 8392: 63; 8674: 60b – De Vogel & Vermeulen 6518: 21; 6608: 21; 7567: 115; 7612: 58; 7617: 100 – De Voogd 118: 101; 220: 6a; 3147: 49a – De Wilde & De Wilde-Duyfjes 12014: 117; 12014A: 56; 12343: 11; 12346: 94; 12626: 6a; 13034: 115; 13225: 121; 13771: 115; 14518: 37; 14604: 102; 14627: 37; 14635: 56; 14723: 115; 14760: 103; 14766: 126; 15219: 110; 15487: 117; 15503: 6a; 15742: 56; 15794: 115; 15864: 56; 16596: 102; 16785: 121; 16786: 110; 18104: 56; 18625: 126; 18720: 102; 18748: 56; 18784: 127; 18816: 29; 18849: 117; 19004: 58; 19729: 102; 19952: 102; 20044: 37; 20231: 40; 20236: 6a; 20299: 117; 20332: 40; 20412: 40; 21238: 29 – Dennis RSS2145: 114 – Dilmy 434: 115 – Din & Shi 232: 49a – Djamhari 68: 101 – Docters van Leeuwen 3251: 127; 3927: 101; 3928: 115; 9200: 25; 10474: 26; 10710: 25; 10727: 25; 10996: 49a – Dorgelo 125: 22 – Dransfield 968: 115; 2487: 64; 6343: 88; 6537: 77; 6854: 72b; 7069: 12 – Dyg Awa et al. 44116: 67. E.G.1532: 123 – E. Hainan Expedit. 530: 89 – Eberhardt 1981: 35; 2050: 49a; 3852: 59 – Edaño PNH21: 11; PNH6617: 11; PNH14225: 20 – Elbert 253: 60a; 254: 73; 4429: 128 – Elmer 5778: 6a; 6977: 6a; 7050: 128; 7597: 85a; 7634: 85a; 7748: 85a; 7787: 73; 8839: 49a; 9536: 73; 9586: 11; 9687: 106; 11297: 117; 11341: 49a; 12148a: 56; 13252: 90; 13613: 26; 13790: 49b; 14007: 128; 14041: 6a; 14153: 56; 14261: 120; 15705: 37; 16522: 128; 16532: 26; 20147: 103; 20520: 117; 20913: 103; 21046: 53a; 21169: 102; 21212: 12; 21358: 29; 21423: 6a; 21634: 53a; 21656: 12; 21865: 30; 21967: 128; 22045: 49a; 22079: 6a – Endert 2661: 76; 2769: 37; 2828: 121; 2973: 117; 3627: 53a; 4096: 37; 4358: 103; 4572: 103; 4814: 54 – Eyma 15: 13; 1954: 13; 4827: 124. Fallen 239: 23 – Fenix 12619: 26 – Forbes 203: 26; 266: 56; 296: 58; 313: 113; 570: 58; 741: 19; 786: 64; 1007: 26; 1074: 64; 1085: 60a; 1817: 101; 2194: 58; 2418a: 75; 2573: 101; 3096a: 37; 3175a: 102; 3707: 128 – Ford 66: 49a; 113: 125a – Fosberg 36993: 128; 37158: 128; 37201: 6a; 37241: 56; 37364: 56; 44588: 125; 44589: 73 – Franken & Roos 306: 127 – Frodin & Pullen UPNG867: 26 – Fuchs 21320: 72b – Fuentes et al. PPI37061: 21; PPI37225: 21; PPI38961: 128; PPI38966: 56 – Fukuoka T-36217: 23; T-62359: 60a; T-62361: 6a ; T-62491: 6a; T-63785: 60a; T-63786: 35; T-63815: 20 – Fukuoka et al. T-177: 6a; T-34554: 60a; T-34620: 60a; T-35953: 117; T-35954: 23; T-35955: 102. Abbe et al. 10030: 20 – Adelbert 470: 64 – Afriastini 1359: 29; 2369: 56 – Afriastini et al. J-67: 58 – Alleizette 3295: 20 – Alston 12951: 58; 14171: 9 – Ambriansyah & Arifin AA351: 12; W1022: 53a – Amdjah 19: 76; 366: 37; 614: 78; 1004: 22; 1006: 12; 1020: 12; 1075: 12 – Anang 238: 56 – D. Andau 1049: 70 – Anderson 88: 6b; 202: 115; 8382: 53b – Andong & Kostermans 23: 35 – Arbain DA239: 125a; DA250: 123a; DA399: 61 – Argent et al. 25383: 102; 25384: 117; 91103: 117 – Arsin 9: 101; 126: 19; 19674: 19 – Atkins et al. 501: 6a – Atmodjo 293: 58 – Avé 24: 123; 99: 58; 100: 123. Backer 3186: 73; 4915: 26; 6058: 56; 6060: 106; 6074: 37; 8957: 117; 9218: 103; 9357: 11; 9929: 11; 9983: 56; 10533: 58; 10699: 103; 10729: 101; 10853: 106; 11063: 102; 13644: 103; 14064: 56; 14163: 73; 14334: 56; 14458: 73; 14599: 19; 14639: 73; 16276: 37; 22515: 23; 22737: 13; 22740: 56; 22988: 101; 23067: 56; 23831: 101; 23934: 56; 25457: 37; 25544: 37; 26272: 89; 31192: 56; 31224: 89; 37545: 73; 37680: 73 – Bakhuizen van den Brink 1555: 64; 1696: 64; 2451: 26; 2472: 26; 2491: 56; 2829: 64; 3673: 26; 3999: 73; 4135: 73; 4337: 11; 4394: 56; 4441: 115; 4456: 11; 4466: 58; 4524: 101; 5539: 101; 7077: 37; 7096: 6a – Balakrishnan 137: 128; 654: 125a; 755: 56; 1664: 128; 3496: 56 – Balansa 640: 20; 2654: 112; 2655: 112; 2656: 6a; 2658: 35 – Barbon et al. PPI8199: 56; PPI8258: 73; PPI9303: 56; PPI12316: 128 – Bartlett 325: 103; 6452: 40; 6552: 47; 6962: 127 – Beaman 7192: 42; 7194: 109; 7244: 103; 7260: 78; 7261: 26; 7276: 59; 7277: 63; 7329: 78; 7368: 103; 7371: 103; 7396: 53a; 7402: 109; 7423: 53a; 7439: 109; 7574: 102; 7643: 102; 7645: 94; 7652: 102; 7656: 37; 7666: 103; 7775: 78; 7783: 70; 7794: 29; 7825: 3; 7837: 102; 8111: 103; 8149: 84; 8170: 63; 8198: 103; 8204: 60c; 8209: 60c; 8225: 103; 8226: 22; 8258: 60c; 8436: 72b; 8444: 11; 8695: 63; 8751: 60c; 8829: 63; 8928: 92; 9088: 42; 9248: 78; 9330: 78; 9337: 103; 9478: 78; 9552: 42; 9828: 12; 9848: 108; 10038: 88; 10163: 37; 10395: 103; 10421: 26; 10579: 63; 10633: 103; 10752: 109; 10788: 103 – Beccari 143: 115; 165: 103; 236: 115; 952: 29; 966: 56; 969: 54 – Berger 531: 19 – Beumée A151: 56; A304: 56; A383: 37; 6713: 29 – Bhargava 3767: 128; 4339: 125a – Bicknell 710: 6a; 860: 26; 942: 56; 964: 85a; 1049: 56 – Blackwood 249: 122 – Bloembergen 3572: 128; 3699: 48; 3906: 122; 3927: 49a; 3939: 103 – Blume s.n. barcode L0000688: 37; barcode L0000691: 64; barcode L0000692: 64; barcode L0000693: 64; barcode L0000694: 64; barcode L0000697: 73; barcode L0000700: 102; barcode L0000701: 102; barcode L0000702: 102; barcode L0000703: 103; barcode L0000707: 115; barcode L0000708: 117; barcode L0000709: 117; barcode L0000711: 127; barcode L0000722: 11; barcode L0057519: 101; barcode L0057520: 128; barcode L0057521: 124; barcode L0057522: 66; barcode L0057523: 66; barcode L0057524: 66; barcode L0057525: 66; barcode L0057526: 66; barcode L0057529: 89; barcode L0305211: 62; barcode L0305213: 62; barcode L0305214: 62; barcode L0305289: 73; barcode L0305583: 115; barcode L0305599: 115; barcode L0310036: 19; barcode L0310418: 54; barcode L0310419: 54; barcode L0310420: 54 – Bogor Botanical Garden 136: 56 – Borssum 1864: 37; 2762: 115; 2804: 122 – Brass 7222: 37; 7223: 25; 12811: 122; 13025: 122; 13639: 26; 13670: 36; 19330: 21; 19883: 21; 25023: 26; 27979: 56; 29258: 21; 29476: 69; 31711: 26; 31765: 26 – Bremer et al.1650: 67 – Brinkman 822: 125a – Britton 173: 117; 374: 11 – Brooke 8480: 102; 8484: 102; 8588: 53a; 9115: 111; 9198: 53a; 9337: 102; 9348: 71; 9708: 22; 10420: 63; 10423: 103; 10735: 127 – Bruggeman 236: 106; 662: 73 – BRUN 143: 121; 559: 22; 930: 12; 5767: 102; 15246: 29; 15640: 46; 15797: 17 – BS series 24565: 128; 39562: 125a; 7405: 6a – BSIP 7668: 114; 8509: 114; 10088: 114; 11253: 21; 11660: 114 – Bubung Sk 301: 130 – Bunchuai 55: 73; 591: 19; 600: 23; 608: 63 – Bünnemeijer 380: 37; 438: 37; 1018: 37; 1448: 6b; 1791: 53b; 2075: 56; 2217: 37; 2498: 56; 2647: 101; 2675: 115; 2677: 103; 2883: 115; 3345: 58; 3578: 61; 3686: 61; 3862: 61; 3946: 103; 5483: 103; 5746: 83; 10335: 9; 11010: 124 – Bur. Sci. 38795: 112 – Burkill 244: 4; 976: 125a; 2064: 107; 2305: 72a; 2844: 3 – Burley et al. 858: 71; 1439: 127; 1700: 127; 2353: 117; 2423: 117; 2741: 104; 2894: 76; 3327: 56 – Burut 3000: 78 – Buwalda 221: 6a; 3181: 89; 3320: 124b; 3477: 115; 3478: 64; 3642: 99; 4713: 48; 6269: 72a; 6528: 127; 6546: 79; 7590: 117; Zhu H. 2000a. A new species of Lasianthus (Rubiaceae) from Peninsular Malaysia. Garden’s Bulletin Singapore 52: 257–259. Zhu H. 2000b. New plants of the genus Lasianthus (Rubiaceae) from Vietnam. Acta Botanica Yunnanica 22: 398. Zhu H. 2001a. New plants of Lasianthus Jack (Rubiaceae) from Kinabalu, Borneo and its biogeographical implication. Blumea 46: 447–455. Zhu H. 2001b. A taxonomic revision of the genus Lasianthus Jack (Rubiaceae) from Thailand. Acta Phytotaxonomica Sinica 39: 116–150. Zhu H. 2002. A revision of the genus Lasianthus (Rubiaceae) from China. Systematics and Geography of Plants 72: 63–110. IDENTIFICATION LIST Numbers after the specimen numbers refer to the species numbers in this revision. 99 H. Zhu et al.: Revision of Malesian Lasianthus Gaerlan et al. 3458: 6a; PPI2655: 49a; PPI19900: 128; PPI23712: 16 – Galoengi 334: 54 – Garcia et al. PPI15287: 56; PPI15290: 21; PPI15979: 85a; PPI18144: 56 – Gardette E.G. 1532: 123; 1914: 6a – Garrett 384: 60a; 879: 60a – Geesink 9034: 103; 9144: 63; 9159: 103; 9271: 60c ; 9326: 117 – Geesink et al. 5035: 23; 5695: 128; 6259: 29; 6493: 56; 6494: 23; 6574: 56; 6601: 29; 6653: 56; 6877: 128; 7374: 23; 7573: 117; 7578: 102; 7755: 56; 7782: 35 – Gjellerup 485: 26; 1228: 73 – Grashoff 680: 127 – Gressitt 292: 49a – Griffith 2929: 73. Halle 1997: 102 – Hallier 26: 81; 67: 115; 120: 64; 164: 106; 200: 101; 232: 117; 245: 64; 333a: 115; 385: 115; 421: 73; 461: 117; 1737: 103; 1738: 115; 1762: 6a; 1919: 81; 1967: 81; 2004: 29; 2016: 71; 2416: 37; 2882: 53a; 2884: 29; 2904: 37; 2981: 22; 3083: 76; 3289: 76 – Hance 18438: 6a – Hansen 295: 72b; 607: 6a; 1622: 111 – Hansen et al. 11088: 6a – Hardial 600: 23 – Harmand 916: 35; s.n. (Pierre Herb. no. 6173): 125a – Hartley TGH10693: 26; TGH10742: 21; TGH10791: 21; TGH11377: 56; TGH12643: 23 – Hasskarl 18121: 6a – Hatusima 18788: 49a; 19169: 49a; 19171: 6a; 20190: 35 – Hatusima & Sako 21542: 49a; 23872: 6a – Haviland 1119: 42; 1141:108; 1241: 78; 1300: 78; 1306: 63; 1323: 63 – Haviland & Hose 3441: 12 – Henderson 18008: 75; 19732: 79 – Hennipman 5214: 26; 6118: 21 – Hochreutiner 222: 62; 241: 101; 249: 101; 250: 64; 781: 29; 1172: 60a – Holstvoogd 353a: 124 – Hoogerwerf 53: 89; 218a: 125b; 237: 89; 250: 124; 252: 59 – Hooker & Thomson 11: 112 – Hoop 51: 37 – Hopkins & Naoni 455: 23 – Hose 117: 12; 156: 123; 634: 68 – Hosseus 353: 73 – Hotta et al. 10: 115; 19: 115 ; 79: 58; 178: 42; 201: 57 – Hou 278: 109; 626: 56; 659: 56 – Hu & But 20974: 20; 21958: 20 – Huang T.C. 6475: 49a – Huang T.C. & Kao M.T. 5134: 56; 6282: 128. Iboet 35: 54; 42: 128 – Idjan 287: 26 – Ijiri & Niimura 694: 21 – Irawati 209: 6a – Ismael 3: 18; 5: 100 – Iwatsuki et al. 17: 49a; P854: 73; 910: 11; P1185: 6a; P1279: 73; P1292: 26; P1373: 73; P1375: 73; S926: 56; S958: 121; T8307: 49a; T8308: 6a. Jaag 363: 21 – Jacobs 4480: 26; 4666: 83; 4824: 128; 7576: 49a; 7796: 85a; 8139: 11; 8158: 59; 8442: 104; 8705: 69 – Jaheri 265: 37; 318: 117; 391: 117; 880: 37 – Jarvie et al. 5218: 53a; 5700: 29; 6063: 102 – Jayasuriya & Premadasa 1647: 128 – Jeng 512: 6a – Johansson et al. 60: 117; 118: 125a; 209: 125a – Johns et al. 129: 124 – Junghuhn 287: 19; 321: 101; 342: 115. Kadir & Enggoh 10339: 72b – Kajewski 2612: 114 – Kalkman 4154: 49a; 4234: 49a – Kamarudin FRI34510: 52 – Kao M.T. 5555: 128; 5939: 49a; 5970: 35; 6678: 49a – Kartawinata 705: 76; 767: 94; 774: 29; 1013: 117; 1029: 102; 1037: 104; 1149: 64 – Kasik 5: 19 – Kasim et al. 01708: 103 – Kato & Miki 273: 35 – Keng H. 1043: 128 – KEP-FRI series 343: 94; 1931: 103; 1957: 29; 1958: 123; 4437: 104; 4881: 131; 5476: 79; 6111: 56; 6443: 79; 7159: 94; 8536: 97a; 10854: 72a; 10973: 14; 11081: 53a; 12325: 6b; 12392: 74; 12655: 20; 12904: 103; 13202: 6b; 13429: 102; 14793: 97a; 15573: 93; 15583: 3; 15716: 34; 16651: 53a; 16861: 53a; 18077: 118; 20188: 53a; 20679: 131; 22027: 41; 23067: 98; 23311: 104; 25173: 97a; 25703: 123; 28310: 53a; 30929: 53b; 32032: 107; 32036: 115; 32705: 103; 35229: 75; 46934: 118; 94488: 87; 98994: 53b; 99775: 94; 104440: 97a; 104635: 72a; 104817: 8 – Kern 7458: 19; 8577: 11; 8618: 56 – Kerr 9845: 6a; 10833: 59; 11941: 97b; 14117: 118; 14940: 29; 15435: 29; 15930: 118; 16700: 97b; 18809: 40; 19424: 11; 21327: 4; 21510: 128; 21591: 128; 21708: 11 – Kessler PK32: 43; PK150: 22 – Kew Dist. 869 (Griffith s.n.): 97a; 871 (Maingay 1699): 102; 2918 (Griffith s.n.): 97a; 2922 (Griffith s.n.): 131; 2929 (Griffith s.n.): 127; 2934 (Griffith s.n.): 40; 2935 (Griffith s.n.): 53a – Kiah 24312: 50 – Kiapranis LAE 69008: 91 – King 857: 45 – King’s collector 237: 6a; 402: 87; 472: 37; 3209: 103; 4128: 87; 4462: 87; 10100: 127 – D.W. Kirkup & S. Thomas 619: 111 – KL series 437: 6a; 489: 87; 1500: 87; 1755: 29; 3492: 103; 3633: 53b – Klackenberg & Lundin 800: 103 – Kleinhoonte 30: 101; 31: 115; 65: 13; 172: 19 – Koelz 27222: 73; 28218: 73; 30127: 60a – Kokawa & Hotta 389: 78; 401: 37; 840: 117; 2218: 103; 2394: 53a; 2843: 109; 4236: 60b; 4991: 37; 4995: 53a; 5382: 53a; 5624: 109; 5916: 53a; 6057: 63 – Koorders 6547: 101; 6549: 64; 21800: 115; 22224: 115; 23209: 60a; 24328: 56; 24335: 64; 24340: 101; 24475: 37; 25782: 115; 26369: 26; 26418: 101; 26456: 64; 26614: 49a; 26710: 60a; 27423: 6a; 31949: 124; 32065: 106; 32858: 64; 32909: 56; 33234: 56; 34814: 60a; 36715: 73; 39097: 115; 39250: 6a; 39513: 56; 40638: 64; 42109: 73; 43989: 29; 44074: 29; 44093: 56; 47940: 6a – Kostermans 4546: 29; 4550: 12; 5720: 12; 6255: 128; 6299: 89; 6305: 56; 6393: 12; 6661: 6a; 7906: 56; 9323: 12; 9872: 29; 10039: 53a; 10493: 76; 10627: 12; 13092: 53a; 13819: 117; 18718: 6a; 21875: 102; 22002: 29; 22011: 102; 22046: 29; 23508: 103; 23916: 64 – Kostermans & Anta 92: 35; 132: 29; 186: 104; 683: 56; 707: 53b; 1297: 53b; 1366: 6b; 1387: 6b – Kostermans et al. 395: 21; 396: 21; 416: 122a; 490: 6a; 521: 21; 653: 26 – Koyama et al. 10591: 123; T31581: 56; T61218: 49a – Krukoff 4061: 103; 4068: 56 – Kuntze 4565: 101 – Kuo C.C. 1172: 6a; 1182: 35 – Kwangtung 786138: 6a; 79-6119: 20; 80-6279: 49a; 82-5696: 125a. LAE series 50043: 21; 51237: 21; 51745: 113; 56802: 21; 57478: 23; 58086: 122; 58297: 49a; 58299: 122; 58400: 26; 58663: 21; 63325: 85a; 64227: 21; 67926: 26; 73543: 21; 73750: 113; 74795: 21; 74813: 21; 77186: 26; 77522: 26 – Lam 19: 103; 154: 115; 176: 19; 309: 26; 310: 73; 793: 21; 797: 25; 936: 25; 975: 25; 1217: 21; 2252: 56; 7708: 21 – Laman et al. TL44: 53a; TL62: 102; TL228: 127; TL232: 37; TL285: 37; TL801: 5; TL990: 117 – Lamont 325A: 6a – Lanjouw 50: 23; 202: 64 – Larsen et al. 198: 23; 808: 6a; 9157: 125a; 32752: 29; 32851: 29; 33176: 40; 33318: 102; 33320: 23; 33461: 97b; 33863: 128; 45393: 125b; 45620: 40; 45955: 27; 46093: 4; 46316: 112; 46436: 89 – Latiff et al. 3948: 33 – Lau S.K. 20212: 20 – Laumonier TFB59: 101; TFB809: 127; TFB1047: 56; TFB3838: 94; TFB3869: 25 – Ledermann 6630: 21; 9373b: 69; 9764: 25 – Lei 1025: 125b – Leonardo 3271: 49a – Lesger 379: 56 – Lörzing 1828: 101; 4719: 58; 4929: 26; 4933: 103; 5243: 56; 5285: 117; 5473: 56; 5606: 81; 5790: 54; 5801: 37; 5886: 54; 5977: 130; 6916: 54; 6972: 56; 8622: 115; 11395: 54; 13729: 103; 14599: 29; 14609: 117; 14929: 130; 15111: 106; 15201: 40; 15915: 56; 16099: 29; 16227: 47; 16434: 56; 16691: 56 – Lűtjeharms 3759: 125a; 4109: 128; 4700: 117. Mabberley 1563: 128 – Madulid 6821: 6a – Madulid et al. PPI3085: 85a; PPI3089: 49a; PPI20187: 21; PPI21048: 128 – Maingay 866: 127; 867: 72a; 868: 6b; 869: 53a; 870: 56 – Manner & Street 519: 49a – Maradjo 378: 54 – Martin S39288: 115 – Maskuri 26: 56; 884: 102 – Mathew S.P. 21144: 6a – Matthew K.M. 26773: 128; 28243: 128 – Maxwell 74-883: 56; 74-924: 11; 75-192: 128; 75-484: 128; 75-793: 59; 76-145: 6a; 76-211: 23; 78-70: 29; 81-48: 104; 82-72: 6b; 82-100: 29; 82-124: 53a; 82-276: 100; 82-279: 53a; 85-1112: 29; 85-1113: 117; 86-30: 20; 87-251: 56; 87-525: 56; 87-710: 23; 87-1040: 23; 87-1194: 6a; 88-578: 89; 90-418: 73; 90-564: 23; 90-1279: 89; 92-195: 49a; 92-332: 23; 93-523: 23; 93-1207: 6a; 94-1235: 125a; 96-1424: 6a; 96-1655: 73 – McDonald & Afriastini 3440: 37; 3441: 102; 3377: 56 – McGregor 32292: 85a – McKee 1861: 23 – Meijer 45: 64; 431: 103; 1006: 12; 1353: 101; 1433: 101; 2433: 37; 2706: 64; 5004: 101; 6151: 9; 6767: 102; 7613: 26 – Merrill 5739: 49a; 7041: 73 – Meyer Drees 41: 111 – Milliken 954: 115; 964: 122 – Mitsuta T45322: 60a; T48760: 73 – Mogea 2347: 73 – Mogea et al. 3590: 22; 3748: 102; 4019: 7; 4020: 53a; 4094: 53a; 4166: 29; 4193: 53a; S52750: 102 – Mohtar et al. S49487: 72b ; S65895: 53a – Moi W. et al. 105: 26 – Monchy de 107: 101 – Mondi 130: 56; 292: 22 – Moulton 2019: 53b – Mousset 148: 113 – Muchtar 46A: 127 – Murata et al. B377: 37; B379: 37; B1220: 53a; B1224: 53a; B1233: 53a; B1724: 22; B1745: 53a; B2301: 12; B2306: 111; B2363: 53a; B2836: 22; B3922: 11; J933: 100; J1914: 115; T15785: 73; T16246: 56; T16248: 23; T16252: 89; T16350: 89; T17438: 29; T17441: 56; T17449: 89; T17596: 23; T37192: 128; T41659: 20; T41672: 23; T42684: 35; T42686: 73; T42850: 60a; T42855: 35; T42856: 35; T42858: 23; T42896: 60a; T51489: 20. Nagamasu 3046: 115; 3098: 101; 3114: 15; 3121: 75; 3140: 57; 3271: 102; 3277: 123; 3359: 15; 3371: 58; 3373: 115; 3390: 75; 3526: 101; 3656: 42; 3678: 115; 3691: 43; 3821: 15; 3836: 75; 3873: 75; 3874: 115; 3979: 42; T49936: 49a – Nair 840: 125a – Nakaike 633: 6a – Nedi 9: 13; 582: 117 – NGF series 8384: 122; 11909: 36; 14310: 21; 14942: 122; 17907: 6a; 19142: 21; 19155: 6a; 19477: 21; 20351: 49a; 20757: 122; 21601: 124; 23419: 124; 25710: 36; 26121: 23; 26665: 21; 28869: 36; 28889: 26; 29101: 122; 31009: 26; 32267: 21; 33453: 117; 33458: 56; 33463: 25; 33469: 117; 33790: 49a; 38035: 117; 39913: 26; 39936: 85b; 41478: 21; 45378: 122; 47659: 49a; 48185: 21 – I. Nielsen & H. Balslev 1086: 111 – Nitrasirirak 215: 102 – Nitta 15124: 115; 15186: 60a – Noguchi 3186: 6a; 3187: 35; 3188: 49a; 3516: 49a – Noor MN48: 125a; SRMN28: 6b – Nooteboom 962: 29; 1077: 37; 1167: 77; 1367: 115; 1450: 53a; 4189: 7; 4203: 76; 4225: 22; 4340: 76; 4560: 117; 4657: 102; 4735: 7; 4743: 102; 5189: 113. Odashima 11131: 20 – Ogata 10618: 103; 10815: 53a; 10865: 102; 11285: 60c; 11297: 103; 11310: 60c; 11567: 37. Panigrahi 3613: 60a; 4136: 73; 4845: 60a – Partomihardjo 1581: 73 – Paul Chai 34805: 51 – Pereira et al. 84: 60c; 90: 60c; 125: 53a; 163: 103; 169: 103; 173: 103; 711: 103 – Phengkhlai 136: 102; 137: 35; 150: 102; 212: 128; 503: 6a; 3234: 20 – Pickles 2934: 115; 3673: 99 – Pierre 587: 128; 1316: 23; 3256: 56; 3257: 20; 3258: 20; 3266: 49a; 3267: 23 – Pipoly et al. 170: 12; PPI38153: 56; PPI38172: 56 – Pleyte 16: 115; 62: 101; 68: 101; 175: 48 – Ploem 18966: 19 – PNH series 995: 73; 1370: 73; 1477: 73; 1910: 117; 2134: 73; 2851: 73; 3260: 73; 3476: 117; 3506: 117; 6095: 6a; 6379: 56; 6606: 56; 6615: 26; 6635: 106; 6677: 26; 6678: 6a; 7127: 85a; 7178: 85a; 7184: 6a; 7197: 85a; 7312: 21; 7378: 85a; 9978: 73; 13747: 117; 14208: 56; 14225: 20; 14487: 6a; 17697: 21; 19771: 6a; 21490: 56; 21496: 6a; 21867: 21; 21875: 128; 21913: 21; 21920: 49a; 21924: 49a; 21984: 73; 22016: 49a; 36122: 6a; 37840: 128; 38481: 21; 38795: 112; 41894: 6a; 72623: 49a; 72624: 85a; 77992: 128; 78175: 6a; 78255: 56; 117173: 128; 117189: 56; 117396: 6a; 117792: 6a; 118056: 128; 118439: 6a – Poilane 234: 60a; 283: 20; 2327: 20; 40832: 23 – Popta 1507: 64 – Posthumus 776: 53a – Prance et al. 30620: 17 – Prawiroatmodjo & Maskuri 1423: 102 – Provencher T56: 6a – Puasa-Angian 3967: 119 – Puff et al. 900808-1/4: 111; 900809-1/1: 117; 900809-1/3: 12; 900812-1/1: 72b; 900818-1/1: 37; 900818-1/8: 117; 900819-1/2: 78; 900829-1/2: 12; 920420-1/3: 37 – Pullen 943: 21; 7424: 21; 8271: 26 – Purseglove P4172: 103; P4518: 40; P4553: 29; P4669: 94; 100 Blumea – Volume 57 / 1, 2012 P4687: 37; P4784: 53a; P4869: 22; P4915: 22; P4931: 53b; P5121: 102; P5218: 102; P5271: 53a; P5335: 117; P5467: 117 – Put 1195: 32. Raap 148: 6a; 166: 64; 200: 101; 241: 115; 252: 26; 538: 29; 564: 81; 743: 115; 778: 115 – Rachman et al. 3: 6b; 4: 56 – Rahayu & Maskuri 381: 125a – Rahmat si Boeea 3484: 29; 5437: 29; 5705: 127; 5757: 127; 6253: 102; 8604: 127; 9232: 81; 9406: 37; 9499: 53a; 9992: 37; 10183: 130; 10185: 103; 10190: 103; 10397: 75; 10582: 75; 10729: 103; 10877: 42; 11032: 130; 11083: 103 – Ramlan 091: 115 – Ramos BSN 13982: 56; 14508: 11; 14619: 85a; 15274: 6a; 41250: 56 – Ramos & Edaňo 30682: 85a; 30865: 56; 38058: 1; 38763: 56 – Ramsri 70: 50 – Rao 9082: 6a – Rao et al. K8015: 103 – Rau K. 389: 2 – Reksodihardjo 137: 64; 140: 102 – Rena et al. S58966: 67 – Reynoso & Majaducon 24890: 11 – Reynoso et al. PPI1157: 21; PPI7333: 128; PPI11170: 128; PPI11373: 128; PPI14273: 21; PPI14448: 85a; PPI14515: 85a – Richards 1010: 12; 1061: 37; 1114: 51; 1284: 102; 1351: 102; 1467: 51 – Ridley 140: 22; 2904: 11; 3219: 74; 6463: 53b; 8213: 37; 8578: 45; 9095: 104; 9703: 12; 9715: 37; 10948: 6a; 11486 (14490 in publication): 52; 12542: 40; 13628: 28; 13630: 82a; 13631: 107; 14007: 93; 14008: 3; 14009: 58; 14487: 33; 14488: 40; 15689: 95; 15690: 98; 16121: 47 – Ridsdale 800: 117; 836: 117; 944: 6a; 983: 6a; 1046: 105; 1152: 117; 1706: 85a; 1711: 85a; 1753: 73; 1757: 85a; 1933: 37; 2456: 37; 2632: 69; PBU365: 117; PBU451: 117; SMHI1701: 128; SMHI1837: 105 – Ridsdale et al. 5476: 11; ISU321: 128 – Robinson 6202: 85a; 6559: 73; 9830: 73 – Romero & Fuentes PPI37505: 128 – RSNB series 215: 111; 250: 37; 798: 108; 839: 108; 1256: 78; 1355: 78; 4033: 109; 4118: 109; 4155: 109; 4598: 109. S series 2936: 117; 3504: 111; 3708: 68; 3873: 116; 4512: 60c; 4513: 63; 8475: 29; 8770: 82b; 12487: 53b; 13619: 39; 14791: 102; 14792: 103; 14793: 115; 16556: 51; 16665: 53b; 16754: 22; 17842: 53b; 18208: 72b; 18876: 39; 19397: 82b; 19719: 53b; 19948: 117; 21641: 12; 21664: 117; 22787: 72b; 22804: 68; 22990: 53a; 23336: 72b; 27239: 51; 28493: 29; 29123: 51; 31102: 29; 31536: 123; 32615: 53a; 32693: 117; 32760: 53a; 32889: 51?; 32936: 55; 33047: 53b; 33331: 76; 33582: 53a; 33809: 10; 33826: 96; 33878: 96; 33966: 103; 34053: 7; 34258: 102; 34913: 53a; 34991: 111; 35169: 39; 35380: 109; 35424: 115; 35572: 103; 36087: 37; 36163: 103; 36226: 53a; 36352: 53a; 36803: 39; 37329: 22; 37447: 127; 39234: 53b; 39325: 53a; 39335: 111; 39588: 72b; 39676: 111; 39988: 99; 40652: 76; 40794: 67; 40861: 53a; 41248: 76; 41928: 59; 41976: 37; 42220: 53b; 42455: 39; 42508: 37; 42843: 111; 43027: 72b; 43146: 72b; 43203: 67; 43843: 29; 44116: 67; 44404: 115; 44480: 60b; 44924: 53a; 45006: 37; 45189: 6a; 45380: 29; 45841: 29; 45874: 53a; 46999: 39; 47207: 51; 47339: 116; 47504: 29; 47533: 6a; 47601: 53a; 47839: 38; 48901: 37; 48941: 116; 49001: 51; 49087: 72b; 49216: 53a; 49382: 53a; 49383: 117; 49399: 53a; 50496: 53b; 50501: 115; 50579: 38; 51624: 103; 51640: 53a; 52056: 53a; 52374: 37; 52556: 31; 52977: 53b; 53651: 53a; 54003: 53a; 54731: 111; 54735: 76; 54758: 12; 56055: 76; 57151: 97a; 57626: 53a; 58317: 102; 58341: 12; 60344: 60c; 61432: 53a; 64427: 12; 66143: 6a – Saito 7046: 35; 7128: 128; 7291: 6a – SAN series 26872: 78; 28270: 78; 33276: 117; 33896: 109; 34923: 72c; 38323: 108; 67607: 12; 69225: 37; 70884: 29; 71057: 102; 74463: 78; 77536: 12; 79334: 12; 79684: 37; 81047: 68; 83017: 37; 83023: 102; 85255: 123; 86828: 56; 90642: 72b; 91035: 94; 93988: 56; 95154: 78; 96680: 109; 97583: 94; 99764: 12; 106675: 109; 107176: 37; 108936: 53a; 111314: 103; 111525: 117; 112143: 94; 115914: 12; 116811: 12; 117593: 12; 118176: 103; 118358: 12; 118556: 46; 118646: 46; 121946: 29; 123429: 53a; 129487: 59c; 132988: 12; 136086: 53a; 136779: 46; 136799: 46; 136864: 111; 141728: 6a – Sands 5244: 60c; 5747: 117; 5848: 117; 5971: 12; 6392: 56; 6482: 56; 6598: 23; 6620: 23 – Sands et al. 1321: 85a; 1469: 21; 1951: 59 – Santos 31887: 49a – Sarip 279: 49a – Sathaphon et al. 104: 117; 247: 37 – Saw FRI36403: 103 – Scheffer 4597: 106; 18929: 56; 18985: 73 – Schiffner 2652: 101; 2660: 73; 2673: 37; 2678: 37; 2679: 56 – Schlechter 17805: 36; 18625: 128; 19334: 37 – Schmidt 683: 102 – Schmutz 56: 6a; 376: 73; 479: 19; 4835: 128; 4867: 48; 5814: 11; 5858: 49a – Schodde 2273: 124; 2897: 21; 2980: 21; 3130: 56 – Scortechini 29: 45; 190: 45; 329: 32; 332: 95; 342: 83; 411: 110; 1207: 37; 6172: 53a – Serian, B. & K. Bremer 1650: 67 – SF series 18962: 78; 32244: 127; 39904: 97a; 40118: 87 – Shah 196: 37; MS1418: 6b; MS1435: 131; MS1540: 97a; P5645: 53b – Shah & Ahmad MS2277: 53a; MS2300: 104 – Shah & Kadim MS507: 97a – Shah & Noor MS658: 103; MS710: 45; MS711: 72a; MS1787: 97a; MS2024: 40 – Shah & Sidek MS1132: 37 – Shea 23884: 117; 26887: 103; 27982: 53a; 28148: 116; 28211: 116; 28267: 81; 28285: 115 – Shimizu et al. M13351: 93; M13352: 103; M13638: 103; M13834: 6b; M13836: 72a; M13836b: 127; M13873: 58; M13962: 104; M13964: 127; T7754: 125b; T8063: 125a; T8858: 20; T14714: 125b; T18113: 60a; T19560: 20; T19583: 89; T19601: 49a; T19603: 6a; T23460: 23; T23729: 56; T23734: 49a; T23782: 49a; T23830: 56; T26200: 97a; T226633: 23 – Simpson 2076: 72b; 2118: 12; 2201: 46; 2451: 102; 2518: 37 – Sinclair et al. 9038: 108; 10439: 53b – Sirirugsa 1034: 4 – Smith J.J. 874: 13 – Smith J.J. & Rant 360: 115; 370: 73; 616: 103; 676: 117 – Smitinand 460: 128; 944: 6a; 1408: 56; 1413: 89; 5689: 56 – Soedarsono 249: 102; 250: 79; 327: 79 – Soegandiredja 83: 26 – Soejarto et al. 6053: 128; 7656: 128; 7807: 128; 8295: 125a; 8312: 128; 8747: 128; 8845: 6a – Soembawa 194: 48 – Soepadmo 160: 29; 908: 118; 1043: 131 – Sørensen et al. 649: 56; 666: 56; 3060: 23; 3061: 23; 6335: 49a; 6339: 49a – Stevens et al. 56: 6a; 364: 37; 370: 12; 416: 102; 489: 70; 524: 117 – Stone 5736: 87; 5985: 98; 7190: 131; 10945: 11; 11053: 98; 11424: 109; 14079: 83; PPI6270: 49a – Stone et al. 522: 85a; 659: 85a; 6178: 128; 8569: 118; 12409: 96; 12518: 104 – Streimann 8460: 85b; 8507: 122 – Stresemann 91: 6a – Sugau et al. JBS178: 12; JBS250: 37; JBS305: 104 – Sundaling SAN135221: 94 – Surbeck 44: 53b – Suvarnakoses 1867: 102. Ta-Yi Liu 6: 49a – Tagawa et al. T472: 49a; T473: 60a; T1214: 60a; T1597: 73; T2859: 73; T5323: 29 – Takeuchi 4754: 25; 5096: 69 – Takeuchi et al. 11397: 113 – Tanaka & Shimada 11124: 128 – Teysmann 1567: 56; 4273: 128; 4540: 56; 8005: 71; 8360: 6a; 10833: 29; 12681: 89; 13787: 6a; 18354: 56; 19330: 56; 19344: 56; 19401: 22; 19414: 56; 19423: 6b; 140216: 22 – Ting & Chen 69: 49a; 74: 6a; 423: 20; 517: 20 – Toxopeus 206: 56; 421: 73 – Tsang W.T. 23940: 112; 29013: 56; 29874: 35; 30333: 20; 30459: 35; 30745: 20 – Tsiang 55: 35 – Tsugaru T61639: 89; T61770: 60a; T61776: 60a – Tukirin 402: 22. Ueda et al. C4350: 49a – Unesco 61: 56; 118: 102; 119: 102 – University of San Carlos 59A: 6a; 553A: 73; 694: 26 – Unknown collector 10410: 11 – UPNG 1325: 113. Valeton 236: 106 – Van Balgooy 3009: 115; 3306: 11; 5800: 37 – Van Balgooy et al. 2839: 101; 2854: 101; 2892: 115; 4289: 101; 5367: 53a; 5396: 5; 5459: 102; 5490: 102 – Van Beusekom et al. 633: 95a; 696: 29; 700: 102; 782: 27; 798: 6a; 1697: 60a; 1822: 89; 1870: 89; 1881: 128; 1885: 118; 2199: 23; 2201: 49a; 2202: 60a; 2675: 6b; 3210: 56; 3210A: 56; 3211: 23; 3231: 50; 3270: 23; 3272: 23; 4116: 128; 4358: 56; 4360: 23; 4452: 89; 4456: 128; 4558: 60a; 4561: 35; 4563: 23; 4564: 49a; 4564A: 49a – Van der Meer-Mohr 34: 56 – Van Kregten VK28: 56 – Van Leeuwen 3203: 117; 3942: 54; 8262: 58 – Van Ooststroom 12924: 115; 13230: 61; 13233: 13; 13956: 101; 13961: 19; 14069: 101 – Van Royen 3183: 56; 3342: 25; 3564: 25; 5153: 56; 5163: 56; 11176: 26 – Van Royen & Sleumer 6356: 25; 6395: 25; 6747: 23; 6870: 23; 7272: 103; 7592: 37; 7686: 37 – Van Slooten 2115: 81; 2251: 102; 2252: 117 – Van Steenis 1114: 29; 1116: 37; 1343: 56; 1361: 37; 2314: 11; 2743: 56; 3738: 75; 3788: 56; 8328: 60a; 9247: 56; 9519: 103; 9523: 103; 9545: 121; 9721: 115; 9886: 126; 9922: 115; 9969: 101; 10819: 115; 12440: 73; 12673: 56; 12772: 60a; 12936: 73; 17542: 101; 20591: 128; 20783: 35 – Veldkamp 7025: 73; 8003: 7; 8004: 53a; 8015: 53a; 8048: 123; 8075: 37; 8144: 76; 8272: 6a; 8474: 22 – Verheijen 704: 6a; 705: 6a; 2359: 48; 2363: 125a; 2365: 128; 2678: 19; 2679: 73; 2933: 19; 2942: 73; 2953: 19; 3015: 73; 3031: 19; 3158: 6a; 3173: 128; 4035: 11; 4073: 73; 4079: 6a; 4760: 11; 4768: 6a; 4769: 19; 4772: 49a; 4781: 49a – Verhoef 90: 102 – Vermeulen 788: 22 – Versteeg 1228: 117; 1406: 56; 1434: 25 – Vidal 390: 85a; 1058A: 128 – Vinas & Lelean LAE59326: 26 – Vreeken-Buijs 11: 72a; 33: 72a. Walker & Tawada 6739: 6a – Walker et al. 5882: 128; 6064: 128; 6642: 128; 6738: 49a; 6975: 49a; 7012: 35 – Wall. Cat. 8306: 106; 8310: 23; 8441 p.p.: 32; 8441 p.p.: 34 – Wang 80524: 23 – Wang C.M. et al. 2249: 49a – Webb et al. CW3500: 56 – Whitford 247: 85a – Whitmore FRI15341: 72a; RSS6051: 114 – Whitmore & Kalima TCW3247: 94; TCW3284: 37 – Whitmore & Sidiyasa TCW3418: 21 – Widjaja EAW4310: 73 – Wilkie 93375: 102 – Winckel 5: 64; 41: 64; 133: 58; 210ß: 13; 1707ß: 115; 1817ß: 101; 1916ß: 100 – Winkler 430: 68; 508: 116; 1364: 86 – Wirawan 70: 102; 132: 40; 174: 102; 228: 115 – Wiriadinata 126: 19; 250: 102; 374: 49a; 3249: 53a; 3324: 29; HW119: 101 – Wiriadinata & Maskuri 617: 102; 637: 37 – Wisse 1024: 19 – Woerjantoro 127: 56 – Womersley & Lam 6233: 21 – Wong 225: 37; 404: 31; 775: 53a; 1569: 72c; 1688: 103; 2275: 70; 2323: 12 – Worthington 13324: 72a – Wray 257: 128; 270: 95; 413: 3; 2019: 117; 2588: 37; 2590: 87; 3451: 37; 3911: 44; 4109: 35; 4128: 94a – Wright 242: 20. Yamazaki 6473: 6a – Yip H.G. 159: 20 – Yoshida 1545: 64; 1605: 115 – Yunnan-expedit. 59-13346: 89 – Yusuf & Wahyono 68: 103; 69: 58. Zahro 108: 56; 114: 115; 115: 11 – Zainudin AZ5469: 6a – Zhu 98-7-01: 60b; 98-7-02: 60c – Zollinger 593: 56; 594: 23; 621: 23; 1277: 64; 1478: 102. 101 H. Zhu et al.: Revision of Malesian Lasianthus Canthium furcatum Miq. (dub. & syn.) Dasus verticillatus Lour. 128 Lasianthus acuminatissimus Merr. 1 acutatus Miq. 2 andamanicus Hook.f. 128 var. ciliatus Deb & M.Gangop. 125a angustifolius King & Gamble 3 apiocarpus (Korth.) Miq. (dub.) appressus Hook.f. 4 atroneurus H.Zhu 5 attenuatus Jack 6 var. attenuatus 6a var. minor H.Zhu 6b var. subsessilis Valeton 6a baasianus H.Zhu 7 barbatus H.Zhu 8 barbellatus Ridl. 125a barbiger Ridl. 9 baviensis (Drake) Pit. 112 bifloroideus H.Zhu 10 biflorus (Blume) M.Gangop. & Chakrab. 11 bordeni Elmer 6a borneensis Merr. 12 brachyphyllus K.Schum. 6a bracteatus Wight 56 bracteolatus Miq. 13 bractescens Ridl. 37 var. rosulatus Ridl. 37 braunianus (Warb. ex Boerl.) M.Gangop. & Chakrab. (excl.) brevidens Craib 59 brevipedicellatus H.Zhu 14 brevipedunculatus H.Zhu 15 brevipes Valeton (dub.) brochidodromus H.Zhu 16 bruneensis H.Zhu 17 bulusanensis Elmer 26 bunzanensis Shimizu 59 cailinianus H.Zhu 18 caloneurus K.Schum. 102 canephoroides Wernham 26 capitatus Blume 19 subsp. pendulus (Ridl.) H.Zhu 93 caudiculatus Ridl. (dub.) chinensis (Champ.) Benth. 20 var. odajimae (Masam.) Kaneh. 20 chinensis auct. 128 chlorocarpus K.Schum. 21 var. curvinervis Valeton 21 chryseus Ridl. 22 chrysoneurus (Korth.) Miq. 23 chrysophyllus Miq. 24 chrysotrichoides Valeton 25 chrysotrichus Lauterb. 25 cinnamomoides Miq. 102 clementis Merr. 26 condorensis Pierre ex Pit. 35 congesticymus H.Zhu 27 conspicuus Ridl. 28 constrictus Wight 29 var. latifolius Craib 29 copelandi Elmer 6a cordatus Merr. 30 coriaceifolius H.Zhu 31 coriaceus King & Gamble 95 coronatus King & Gamble 32 crassifolius Ridl. 95 crassinervis Ridl. 33 crinitus Hook.f. 34 culionensis Elmer (excl.) cupreus Pierre ex Pit. 125a curtisii King & Gamble 35 cyaneus Merr. (excl.) Lasianthus (cont.) cyanocarpoides Valeton 36 cyanocarpus Jack 37 var. asperatus Pierre ex Pit. 56 var. lucidus Pierre ex Pit. 56 var. novoguineensis Valeton 56 var. subsessilis King & Gamble 37 cyanocarpus auct. 56 densifolius Miq. 6a, 6b var. calycinus King & Gamble 6a var. latifolia King & Gamble 6a densifolius auct. 6b depressineurus H.Zhu 38 dinghouanus H.Zhu 39 dinhensis Pierre ex Pit. 20 var. glabrescens Pierre ex Pit. 20 var. tonkinensis Pit. 20 ellipticus Wight 40 eriocalyx H.Zhu 41 euneurus Stapf 42 everettii Merr. 56 fasciculus H.Zhu 43 ferrugineus King & Gamble 44 filiformis King & Gamble 45 var. bracteata King & Gamble 45 filipedunculatus H.Zhu 46 firmus Miq. (excl.) flavicans King & Gamble 102 flavinervius Ridl. 47 floresensis H.Zhu 48 fordii Hance 49 var. fordii 49a var. microphyllus (Elmer) H.Zhu 49b formosensis Matsum. var. hirsuta Matsum. 35 foxworthyanus Craib 50 furcatoides H.Zhu 51 furcatus (Miq.) Bremek. (dub.) gainii M.Gangop. & Chakrab. 11 glaber Ridl. 52 glaber Scheff. ex Valeton 101 glabrescens Miq. 23 gracilis King & Gamble 11 griffithii Wight 53 var. griffithii 53a var. latibracteata King & Gamble 53a var. scabridus (King & Gamble) H.Zhu 53b harveyanus King & Gamble (dub.) var. longifolia King & Gamble (dub.) hexandrus (Blume) Blume 54 hiiranensis Hayata 125b hirsutisepalus H.Zhu 55 hirsutus (Roxb.) Merr. 56 hirtimarginatus H.Zhu 57 hirtus Ridl. 58 hispidulus (Drake) Pit. 59 hispidus Elmer (excl.) hoaensis Pierre ex Pit. 23 humilis Elmer 106 inaequalis Blume 37 var. angustibractea Valeton 37 inodorus Blume 60 subsp. inodorus 60a subsp. montigenus H.Zhu 60b subsp. pubescens H.Zhu 60c iteoides Valeton ex H.Zhu 61 iteophyllus Miq. 62 kamputensis Pierre ex Pit. 49a kinabaluensis Stapf 63 kurzii Hook.f. 23 var. fulvus C.Y.Wu & H.Zhu 89 var. howii H.S.Lo 23 var. sylvicola H.S.Lo 89 INDEX Accepted names are in roman type. New names are in bold type; synonyms and superfluous names are in italics. The number after each name is the number of the species as used in this revision; (dub.') refers to dubious taxa, (dub.& syn.) to synonyms of dubious taxa, and (excl.) to excluded from Lasianthus. Lasianthus (cont.) kurzii auct. 22 kwangtungensis Merr. 35 laevicaulis Kurz 56 laevigatus Blume 64 lancilobus H.Zhu 65 langkokensis (Drake) Pit. 112 latifolius (Blume ex DC.) Miq. 66 laxifloroideus H.Zhu 67 laxiflorus Merr. 68 ledermannii Valeton 69 lei Merr. & Metc. ex H.S.Lo 125b leucocarpus (K.Schum. & Lauterb.) M.Gangop. & Chakrab. (excl.) linearifolius H.Zhu 70 longibracteatus H.Zhu 71 longifolius Wight 72 subsp. laevis H.Zhu 72b subsp. longifolius 72a var. obtusifolius H.Zhu 72c lowianus King & Gamble (dub.) lucidus Blume 73 lucidus King & Gamble (dub. & syn.) maingayi Hook.f. 102 var. hirta Ridl. 104 malaccensis King & Gamble 74 malaiense H.Zhu 75 megaphyllus H.Zhu 76 membranaceoideus H.Zhu 77 membranaceus Stapf 78 var. firma Stapf 78 microphyllus Elmer 49b minahassae Koord. (excl.) mindanaensis Merr. 112 minutiflorus H.Zhu 79 mollis Ridl. 80 montanus King & Gamble (dub.) morus Elmer 128 mucronulatus (Korth.) Miq. 81 var. lineata Miq. 115 multibracteatus Wernham (excl.) myrtifolius Ridl. 82 subsp. myrtifolius 82a subsp. vicarius H.Zhu 82b neglectus Valeton ex Bakh.f. 26 nervosus King & Gamble 83 nigrescens H.Zhu 84 nigrocarpus Masam. 128 novoguineensis (Warb. ex Boerl.) M.Gangop. & Chakrab. (excl.) obliquinervis Merr. 85 subsp. novoguineensis H.Zhu 85b subsp. obliquinervis 85a var. taitoensis (Shimizu) T.S.Liu & J.M.Chao 128 obliquinervis auct. 128 oblongatus Merr. 86 oblongus King & Gamble 87 obovatibracteatus H.Zhu 88 obscurus (Blume ex DC.) Miq. 89 oculus-cati Miq. 56 ovatus (Korth.) Miq. (dub.) var. oligoneurus (Korth.) Boerl. (dub. & syn.) palawanensis H.Zhu 90 palembanicus Miq. 81 papuanus Wernham 91 pedicellatus H.Zhu 92 pendulus Ridl. 93 perakensis King & Gamble 94 var. kalimantanensis H.Zhu 94b var. perakensis 94a pergamaceus King & Gamble 95 phymatodeus H.Zhu 96 102 Blumea – Volume 57 / 1, 2012 Lasianthus (cont.) pilosus Wight 97 var. angustifolius Hook.f. 97b var. glabra King & Gamble 97a var. pilosus 97a var. retosus (Wight) Ridl. 97a plagiophyllus Hance 6a platyphyllus (Korth.) Miq. (dub.) poilanei Pit. 60a politus Ridl. 98 var. insularis Craib 32 polycarpus Miq. 53a pseudolongifolius H.Zhu 99 pseudo-lucidus King & Gamble 35 pseudo-stipularis Amshoff ex Bakh.f. 100 pterospermus Wight 127 purpureus Blume 101 reticulatus Blume 102 var. polydasys Miq. 104 retosus Wight 97a retosus auct. 33 rhinocerotis Blume 103 ridleyi King & Gamble 104 ridsdalei H.Zhu 105 rigidus Miq. 106 robinsonii Ridl. 107 robustus King & Gamble 97a rotundatus Stapf 108 roxburghii Wight 56 rufus (Korth.) Miq. (dub.) sabahensis H.Zhu 109 salicifolius Ridl. 3 scabridus King & Gamble 53b scalariformis King & Gamble 110 scalaris Craib 40 sepalinus Ridl. 12 setulosus H.Zhu 111 sikkimensis Hook.f. 112 subsp. langkokensis (Pit.) H.Zhu 112 singalensis Miq. 127 singaporensis King & Gamble 104 sogerensis Wernham 113 solomonensis H.Zhu 114 sordidus Ridl. 40 spathulatus Miq. 53a Lasianthus (cont.) stercorarius Blume 115 var. borneensis Miq. ex Valeton & Winkler 81 var. palembanicus (Miq.) Miq. 81 var. polypleura Blume ex Miq. 115 stercorarius auct. 23 sterrophyllus Merr. 116 stipularis Blume 117 var. hirtus Ridl. (dub.) var. novoguineensis Valeton 117 subaureus Craib 118 subglobosus H.Zhu 119 subinaequalis King & Gamble 37 submembranifolius Elmer 120 subsessilis Miq. 128 subspicatus King & Gamble (excl.) sumatraensis H.Zhu 121 sylvestris Blume 128 var. latifolia Miq. 125b var. subsessilis (Miq.) Miq. 128 sylvestroides Valeton 122 taitoensis Shimizu 128 tamirensis Pierre ex Pit. 128 tashiroi Matsum. 49a tentaculatus Hook.f. 106 tenuifolius H.Zhu 123 tobingensis Miq. (excl.) tomentosus Blume 124 var. hiiranensis (Hayata) H.Zhu 125b tomentosus auct. 125b tonkinensis (Drake) Pit. 6a trichophlebus Hemsl. 125 subsp. barbellatus (Ridl.) H.Zhu 125a var. latifolius (Miq.) H.Zhu 125b var. trichophlebus 125a truncatus Bedd. 128 tsangii Merr. ex Li 112 tubiferus Hook.f. 60a umbellatus Kuntze 101 undulatus H.Zhu 126 velutinus Ridl. 40 venosus Blume 127 venosus auct. 131 venulosus Ridl. 40 verticillatus (Lour.) Merr. 128 var. taitoensis (Shimizu) H.Zhu 128 villosus Ridl. (excl.) Lasianthus (cont.) vriesianus Miq. 129 vulcanicus Ridl. 130 wallichii (Wight & Arn.) Wight 6a subsp. plagiophyllus (Hance) C.Y.Wu & H.Zhu 6a var. penangensis Hook.f. 32 var. setosus (Craib) C.Y.Wu & H.Zhu 6a wightianus Hook.f. 131 wrayi King & Gamble 128 zambalensis Elmer 49a Litosanthes biflora Blume 11 brauniana Warb. (excl.) gracilis Hayata 11 leucocarpa K.Schum. & Lauterb. (excl.) novoguineensis Warb. ex Boer. (excl.) Mephitidia apiocarpa Korth. (dub.& syn.) attenuata (Jack) DC. 6a baviensis Drake 112 var. glabra Korth. 6a capitata (Blume) DC. 19 var. ferruginea Korth. 22 chinensis Champ. 20 chrysoneura Korth. 23 hexandra Blume 54 hispidulus Drake 59 inodara (Blume) DC. 60a langkokensis Drake 112 latifolia Blume ex DC. 66 lucida (Blume) DC. 73 mucronulata Korth. 81 obscura Blume ex DC. 89 odajimae Masam. 20 ovata Korth (dub.& syn.) var. oligoneura Korth. (dub.& syn.) plagiophylla (Hance) Nakai 6a platyphylla Korth. (dub.& syn.) rufa Korth. (dub.& syn.) stipularis Korth. 117 tashiroi (Matsum.) Nakai 49a var. kotoensis Nakai ex Saski 49a tonkinensis Drake 6a wallichii Wight & Arn. 6a Nonatelia? hispida Wall. ex Roxb. 6a Plectronia culionensis Merr. (excl.) Santia Wight & Arn. var. microphyllus (Elmer) H.Zhu 49b Triosteum hirsutum Roxb. 56