A REVISION OF KOPSIA (APOCYNACEAE: RAUVOLFIOIDEAE)
DAVID J. MIDDLETON1
Abstract. The Asian genus Kopsia (Apocynaceae: Rauvolfioideae) is revised. Calpicarpum and Kentrochrosiaare included in synonymy. Approximately 2000 herbarium specimens from 35 major herbaria have been stud-ied. The genus is found from southern China and Burma to northern Australia and Vanuatu and is most diversein Peninsular Malaysia and Borneo. Twenty-three species are recognized, two of which have two varieties each.Three new species are described: Kopsia grandifolia, Kopsia rosea, and Kopsia sumatrana. A key to the speciesis given, all taxa are described with notes on distribution and habitat, distribution maps are provided for all taxa,and all names and synonyms are typified.
This work on Kopsia is the latest in a seriesof revisions of the Asian genera of Apocyn-aceae, subfamilies Rauvolfioideae and Apocy-noideae. The genus contains arguably the mostattractive species of any of the Asian genera ofApocynaceae, and a few of them have becomewidely cultivated. It is possibly also the mostfrustrating genus in the family in Asia for reasonsclarified later. Most of the work for this revi-sion has been done through the study of herbar-ium specimens, but a number of the specieshave also been seen in the field or in cultivation.
Kopsia was published by Blume (1823) inhonor of the Dutch botanist J. Kops,(1765–1849) with one species, K. arboreaBlume. He added another, K. vincaefloraBlume, in his Bijdragen (Blume, 1826) and athird, K. flavida Blume, in Blume (1849). DeCandolle (1844) transferred Cerbera fruticosaRoxb. to Kopsia before the publication of thethird of Blume’s species, and the number ofspecies then grew slowly to the present daywith significant publications of new taxa byHooker (1882), King and Gamble (1907),Ridley (1923), Merrill (1926, 1929), Pitard(1933), Timmerman-Van der Sleesen (1960),Markgraf (1973), Allorge (1993), Allorge andTeo (1986), Middleton (2003), and others. Theonly modern work on the entire genus was a
synopsis of Kopsia in a paper mostly on thechemotaxonomy of the genus (Sévenet et al.,1994). Twenty-six species were recognized inthis paper, and many of the taxa were effec-tively typified (but with extensive applicationof Article 9.8 of the International Code ofBotanical Nomenclature [Greuter et al., 2000]).However, no key was given and many of thetaxa were not described. Other important worksinclude Timmerman-Van der Sleesen (1959) onthe Malesian species, Li et al. (1995) on theChinese species, and Middleton (1999) on theThai species. Kopsia Blume from 1823 is aconserved name against the Orobanchaceaegenus Kopsia Dumort. from 1822.
Calpicarpum was published by Don (1837)with two species, C. roxburghii and C.lamarkii. Unfortunately both species are illegit-imate under Article 52.1 of the Code (Greuteret al., 2000) because Cerbera fruticosa Roxb.was cited in synonymy of Calpicarpum roxburghii, the epithet of which should havebeen taken up, and several species of Cerberawere cited in synonymy of Calpicarpumlamarkii, one of which should have been takenup. De Candolle (1844) effectively typified thegenus with C. roxburghii by excluding the onlyother original species when he synonymizedthe genus under Kopsia.
I would like to thank the Arnold Arboretum and the Harvard University Herbaria, the Nationaal Herbarium Nederland,Leiden Branch, and the Muséum National d’Histoire Naturelle in Paris for their support of this revision. I would also liketo thank the directors and curatorial staff of the herbaria that hosted my visits or loaned material. I am grateful to DavidGoyder for his help in locating specimens at Kew, the Harvard University Herbaria staff for their practical support, KanchiGandhi for fruitful discussions on nomenclatural problems, Holly Nixon for the illustrations, Martin Pullan for help withthe Pandora database, and Carroll Wood and Peter Stevens for helpful comments on the manuscript.
Kentrochrosia was published by Schumannand Lauterbach (1900) with one species, K.monocarpa. Merrill and Perry (1941) sug-gested that all the species of Kopsia withspurred fruits should actually be inKentrochrosia and those with unspurred fruitsshould remain in Kopsia. They acknowledgedthat the genera were indistinguishable onflower characters. If interpreted strictly thiswould leave only Kopsia arborea in Kopsia andall the rest would be moved to Kentrochrosia.Merrill & Perry (1941) made only two newcombinations in Kentrochrosia to add to theoriginal species. Markgraf (1973) lateracknowledged that the single fruit characterwas not only insufficient to distinguish the twogenera but also suggested that the character wasonly one of degree rather than of a qualitativenature. I agree with this interpretation.
The genus Kopsia is most diverse inPeninsular Malaysia and in Sarawak. In bothregions there are a number of species with veryrestricted distributions, and for some of thespecies the collections are few. It is interestingto note that the diversity is not as high in otherparts of Sundaland such as Sumatra orKalimantan, but it should also be recognizedthat these regions are not as intensively col-lected, and one could speculate that more taxawith equally restricted distributions, over andabove the new species K. sumatrana describedbelow, could be discovered when more exten-sive collections are made.
This revision of Kopsia has taken muchlonger than it ought to have for a genus withonly 23 species because of the ease with whichI have been able to set the work aside and dosomething less frustrating. This frustration hasarisen because a large percentage of the avail-able herbarium material, generally speaking,has been very poorly collected. Many of thespecimens have no corollas but rather only the
inflorescences with the ovaries and calyx. Theflowers are extremely delicate, and unless theplant is pressed in the field the corollas will falloff. Even plants pressed immediately oftenmake rather poor specimens, as the corollas arevery thin and delicate and tear and fall off eas-ily. Markgraf (1973) hints at a similar frustra-tion in the introduction to his work on theMalesian species, and he therefore provided akey for plants with “incomplete inflores-cences.” Also, the majority of collections madehave inadequate notes. Only a small percentagenote such characters as tree height and diame-ter or bark color and texture, and few even notecorolla color or whether the “eye” is colored inotherwise white flowers. This “eye” characterwould appear to be a good character for recog-nizing some species such as K. flavida and K.singapurensis, where the “eye” is red in an oth-erwise white corolla. However, it is difficult toknow for sure whether it is consistent in otherspecies with a yellow “eye” such as K. pauci-flora, where the “eye” does not have as much ofa contrast with the white of the corolla, andmost labels mention the color only as whitewith no specific mention of the “eye.” In somespecies like K. arborea, the color is consis-tently white without an “eye.” It will be inter-esting to test this revision against concertedfield observation and better collections in thefuture to see if it will stand the test of time. Inthis regard there are many areas where morecollections are needed, particularly in Vietnam,Kalimantan, and Sumatra.
In the most recent subclassification of theApocynaceae (Endress and Bruyns, 2000)Kopsia is placed in the tribe Vinceae, subfamilyRauvofioideae. This tribe, however, is onlyweakly supported as monophyletic, after theaddition of fruit and seed characters to the mol-ecular data, in an analysis by Potgieter andAlbert (2001).
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Vegetative CharactersHabit. All species of Kopsia are shrubs or
small trees as part of the understory vegetationin forests, at forest edges, or in the open. Thelargest of any species recorded is 14 m tall forK. arborea. However, this is one of the com-monest species, and the heights given in thedescriptions must be judged against the lack ofdata recorded on many specimens, the smallnumber of collections made for several of the
species, and the fact that it is easier to collectsmaller individuals. It is quite probable thatmany of the species have individuals taller thanin the descriptions given.
Bark. Bark characters are very rarelyrecorded on herbarium material so again thecolors given in the descriptions should not beconsidered prescriptive.
Branchlets. Most species of Kopsia haveyoung branchlets that are somewhat angled.
MORPHOLOGY
This may be an artefact of drying in some, butnot all, cases. In some species the anglesbecome so pronounced as to become wing-like.Markgraf (1973) recognized a new species, K.lancifolia, based in part on this character.However, it is somewhat subjective as to whereone draws the line between markedly angledand winged, and all stages in-between are man-ifest in some species, particularly in both vari-eties of K. pauciflora.
Leaves. The leaves are always opposite. Theymay or may not be sessile, a character which issomewhat subjective as many species have veryshort petioles. However, in only two species, K.teoi and K. pauciflora, are there cases wherethe petioles do appear to be absent. In K. pau-ciflora this is of rare occurrence. The bases ofthe petiole weakly clasp the node but withoutthe development of distinct ocrea. Whenyoung, the petiole bases envelop the terminalbud. Small colleters are present in the leaf axils.The blade is always entire and mostly elliptic.The venation is pinnate. The number of veins,the angle of the secondary veins with themidrib, and how distinct the secondary and ter-tiary venation are from each other are usefulcharacters in some species.
Reproductive CharactersInflorescence. Inflorescence structure has
been discussed by Markgraf (1973). The basicstructure is a terminal dichasium or sometimesa trichasium. When the inflorescence is youngthis is the predominant appearance in more orless all species. Some species keep this struc-ture even as they develop and mature (e.g., K.griffithii, K. sleeseniana, K. tenuis, Fig. 6),often becoming somewhat thyrsoid (e.g., K.arborea, Fig. 2). In other species, the dichasialbranches do not further branch and insteadform cincinnate monochasia with flowersopening sequentially one at a time (e.g., K.grandifolia, K. macrophylla, K. pauciflora, K.rajangensis, Fig. 5). In several of the specieswith laxer, more dichasial and open inflores-cences, the ultimate branches are short and theflowers congested (e.g., K. singapurensis, Fig.7). Markgraf (1973) has suggested an evolu-tionary progression from the thyrsoid type in K.arborea to loose di- and trichasia, to di- ortrichasia with cincinnate branches, to cincin-nate branches only. There is a subtending bractat the base of each pedicel and most specieshave further bracts on the pedicels, but this isvariable in some species, and only in K. teoi,with several bracts on the pedicel, is this a use-
ful taxonomic character. Although the inflores-cence is almost always terminal, very rarely arethere also axillary inflorescences (e.g., in K.rajangensis). Sometimes the peduncle is veryshort before the inflorescence branches, givingthe appearance of several inflorescencesemerging from a single apex. In the taxondescriptions the term “inflorescence axes” isused to include the peduncle plus the branchesof an inflorescence.
Calyx. The sepals are mostly ovate, tendingtoward oblong. The apex can be acuminate torounded, and this character is useful for recog-nizing species. All species have a small-to-large gland just below the apex of the sepal.This gland is sometimes decayed or eaten, giv-ing the sepal a secondarily retuse apex. Thereare no colleters in the sepal axils.
Corolla. The basic salverform structure of thecorolla is rather invariable in the genus. Thetube is always slightly dilated around the sta-mens. The corolla lobes are dextrorsely con-torted in bud, a character rather unusual in thesubfamily Rauvolfioideae but invariable withinthe genus. They are held perpendicular to thetube in the open flower and can vary by as muchas 25% in size within an individual flower. Ashas been discussed earlier, the corolla color isoften not recorded on herbarium specimens, butin all known cases the flowers are showy andopen during the day. The appearance wouldsuggest butterfly pollination, although I amunaware of any studies having been conductedin the genus. Useful taxonomic charactersinclude the overall dimensions of the corolla,the color and the shape of the corolla lobes, andthe color of the area where the lobes join thetube, in this work referred to as the “eye.”
Stamens. The stamens, as is the case for mostmembers of subfamily Rauvolfioideae, are ratherunspecialized. The filament is short and theanthers ovate and without lignified guide rails atthe edges. They are attached to the inside of thecorolla tube just above the height of the stylehead. The only taxonomically useful character inthe stamens is their position in the corolla tube.
Disk. There are two disk lobes alternating withthe carpels. In almost all species they are more orless awl-shaped or simply oblong, with an acuteor acuminate apex. In K. arborea they are morevariable within the species and can be some-what more complex in shape (see descriptionbelow). Only in K.griffithii is the disk pubescent.
Gynoecium. The gynoecium is of two ovaries,which are free from each other but are then
united into a common filiform style. The stylehead has a basal collar and a short cylindricalapex. The degree of pubescence, or otherwise, ofthe ovaries is a useful taxonomic character. Eachcarpel contains two ovules, although only oneever develops to maturity. The receptive stig-matic surface is on the sides of the style head.
Fruit. The fruit of all species is a smalldrupe, less than 4.5 cm long, usually pairedalthough sometimes with one aborting, espe-cially in Kopsia arborea. In K. arborea the fruitis ellipsoid or slightly falcate. This is in con-trast to all other species, where the falcate fruithas a spur, or at least a sharp angle, on the ven-tral side. Sévenet et al. (1994) suggest that thereis an evolutionary progression from the fruitwithout an appendage to the large appendagesof K. pauciflora and K. macrophylla. Thelargest appendages, however, are to be found inK. flavida, where it can be up to 13 mm long.No analyses have been done to examine theseevolutionary theories more closely. Markgrafand Huber (1975) give a detailed description ofthe development of the spur in K. flavida. In afew species the fruit is not known, but in thosewhere it is the shape of the fruit and spur is auseful taxonomic character. The color of thefruit is unknown in most species.
Other CharactersThe paper by Sévenet at al. (1994), in which
a synopsis of the genus Kopsia was given, wasprimarily a paper on the chemotaxonomy of the
genus. They noted that alkaloids have longbeen known in the genus and went into somedetail on the chemical structures and rearrange-ments possible. They then presented a tableshowing the alkaloid compounds that havebeen isolated from 15 species of the genus(only 12 recognized in this new revision) andsuggested a number of relationships betweentaxa. I find only a couple of these suggestionslikely. In particular they recognize K. arborea,K. pitardii, and K. jasminiflora as separatespecies, give no suggestions as to the chemicalaffinities of K. arborea but link each of theother two taxa to K. pauciflora and K.dasyrachis, respectively. It is clear to me that K.arborea, K. pitardii, and K. jasminiflora aresynonymous. The suggestions that K. singa-purensis and K. teoi are close, and K.dasyrachis and K. macrophylla are related,would, however, appear to be borne out by themorphology. These issues can be resolved onlyby a more thorough phylogenetic analysis,preferably also using molecular data.
Chromosome numbers are known for K.arborea (n = 36; 2n = 72) and K. fruticosa (n =18; 2n = 36) (see Van der Laan and Arends,1985). In neither the Leeuwenberg classifica-tion system followed by Van der Laan andArends (1985), nor by comparison to theEndress and Bruyns (2000) system, is therebasic chromosome number consistency withinthe tribes in which Kopsia has been included.
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Approximately 2000 herbarium specimensfrom Asia and Malesia have been studied. Theseare from the following herbaria: A, AAU, ABD,AMES, B, BISH, BKF, BM, BO, BR, BRI, C,CAL, CANB, E, F, FU, G, GH, IBSC, K, L, M,MICH, MO, NY, P, PE, PNH, SING, TCD, U,UC,US, Z (Holmgren et al., 1990). All specimenshave been seen unless otherwise indicated.
All dimensions given are for dried specimensexcept for androecium and gynoecium charac-ters, which are from flowers reconstituted by
boiling in water. The number of flowers dis-sected is almost always fewer than the numberof flowers measured for corolla tube and lobedimensions. Therefore, it should be noted thatdimensions given for characters like stameninsertion are based only on the rehydrated flow-ers that were dissected, leading to a few appar-ent discrepancies with the range of variation inthe dried flowers.
Author citation follows Brummitt and Powell(1992).
Shrubs or small trees, buttresses absent.Branchlets terete to winged; glabrous or morerarely pubescent; lenticellate or not. Leavesopposite; mostly petiolate, rarely sessile, basesmostly clasping the terminal bud when youngand the stem when older; blade mostly subcori-aceous to coriaceous, rarely papery, the oppo-site pair more or less equal in size, entire;colleters present in the axils; secondary vena-tion pinnate, clearly distinguishable or not fromthe tertiary venation, often anastomosing intoan intramarginal vein. Inflorescences terminal,very rarely also with some axillary; basicallydichasial, more rarely trichasial, but often withelongated branches that do not further branchso as to appear somewhat cincinnate; peduncu-late or not; glabrous to densely pubescent; sub-tending bracts and pedicel bracts small.Flowers 5-merous. Sepals erect, mostly ovate,sometimes somewhat oblong; without colletorsin the axils; with a gland on the outside justbelow the apex. Corolla actinomorphic; lobesdextrorsely contorted in bud; salverform with a
narrow tube, slightly wider around the stamens,and spreading lobes; tube generally pubescentinside around the anthers and more densely inthe throat, sometimes glabrous in throat, mostlyglabrous outside or rarely with a few hairs onthe lobes and top of tube. Stamens insertedaround the middle of the tube to near the tubethroat, very rarely near base, not exserted fromthroat; filaments straight, short, thin; anthersovate, fertile for most of length; free from stylehead. Disk of two lobes alternating with thetwo free carpels. Gynoecium 2-carpellate,apocarpous but apically united into a commonstyle, glabrous or pubescent; style filiform,glabrous; style head with a collar at the base,otherwise short and cylindrical. Ovules 2 ineach carpel although only 1 ever develops.Fruits of paired drupes, more rarely solitary,ellipsoid to falcate, more or less flattened(hardly at all in K. arborea), usually with spur-like appendages facing inward toward eachother (except in K. arborea). Seed curved,broader at one end, other end acuminate.
rarely less but then corollas not white with a red “eye”. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1616a. Anther apex > 5 mm from corolla throat; inflorescences > 10 cm long, densely puberulent . . . 3. K. dasyrachis16b.Anther apex ≤ 5 mm from corolla throat; inflorescences length variable but mostly < 10 cm long, puberulent
or not . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1717a. Inflorescences with subtending bracts as large as the sepals and flowers clustered at the ends of the inflores-
cence branch; sepals mostly acute, more rarely obtuse; fruit without a spur or angle. . . . . . . . . . . 2. K. arborea17b. Inflorescences with subtending bracts somewhat obscure or smaller than sepals, if as large as sepals then
flowers not clustered at ends of the inflorescence branch; sepals obtuse to rounded; fruit with a spur or angleon one side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18a. Inflorescence axes ≤ 1 cm long; corolla tube < 23 mm long; Hainan . . . . . . . . . . . . . . . . . . . . 9. K. hainanensis18b. Inflorescence axes > 1 cm long, rarely less but then corolla tube > 25 mm long; corolla tube mostly > 23 mm
long, rarely less but then axes > 1 cm long; not in Hainan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1919a. Ovaries glabrous or with just a few isolated hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2019b.Ovaries clearly pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2320a. Leaves with long caudate apex; secondary veins at 40–60˚ to midrib; Borneo . . . . . . . . . . . . 16. K. rajangensis20b.Leaves acuminate; secondary veins at (45–)60–75˚ to midrib; not in Borneo . . . . . . . . . . . . . . . . . . . . . . . . . 2121a. Stamens inserted at < 0.8 of corolla tube length; Vietnam . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. K. tonkinensis21b.Stamens inserted at ≥ 0.8 of corolla tube length; southern Thailand and Malesia . . . . . . . . . . . . . . . . . . . . . . 2222a. Stamen bulge slightly below top of corolla tube; flowers white or pinkish but not white with a strongly
contrasted pink or red “eye”; southern Thailand and peninsular Malaysia . . . . . . . . . . . . . . . . . . . . 17. K. rosea22b.Stamen bulge at top of corolla tube; flowers white with a strongly contrasted pink or red “eye”; Philippines,
eastern Malesia, western Pacific Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. K. flavida23a. Inflorescence robust and lax with distinct internodes all through inflorescence . . . . . . . . . . . 19. K. sleeseniana23b. Inflorescence robust or delicate with flowers congested, if lax then at least with ultimate branches
congested. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2424a. Inflorescences without elongated branches; corolla pale to dark pink or white with a red or pink “eye” . . . . 2524b. Inflorescences with elongated branches but with the flowers congested along these; corolla white or white
with a yellow “eye,” very rarely with a pink tinge but then only ever one flower on an inflorescence branchopen at a time . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
25a. Branchlets pubescent; native in Burma but widely cultivated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. fruticosa25b.Branchlets glabrous; Malesia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2626a. Peduncle to 1 cm long; fruit with a large deltoid or very slightly hooked projection on one side; Philippines,
eastern Malesia, western Pacific Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. K. flavida26b.Peduncle mostly > 1 cm long, very rarely under 1 cm; fruit with only a slight angle on one side; Peninsular
Malaysia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18. K. singapurensis27a. Secondary veins at (55–)60–80˚ to midrib; anther apex 2.4–5.0 mm from corolla throat . . . . . 14. K. pauciflora27b.Secondary veins at 40–60˚ to midrib; anther apex 1.1–2.0 mm from corolla throat . . . . . . . . 16. K. rajangensis
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1. Kopsia angustipetala Kerr, Kew Bull. 1937:42. 1937; Sévenet et al., J. Ethnopharmacol. 41:149. 1994. TYPE: THAILAND. Nong Khai:Chaiyaburi, A.F.G. Kerr 21325 (Lectotype: K,designated by Sévenet et al. [1994];Isolectotypes: BM, P, TCD). Map 1; Fig. 1.
Tree to 5 m tall. Branchlets glabrous todensely puberulent, becoming glabrescentwhen older, sparsely lenticellate or not, weaklyangled. Leaves: petiole 1–7 mm long, glabrous;
blade 3.0–15.8 × 0.9–4.3 cm, 2.4–7.0 times aslong as wide, papery, subcoriaceous to coria-ceous, elliptic, oblong or obovate, apex caudateor long acuminate with a blunt tip, basecuneate, glabrous above and beneath, midribshallowly sunken or raised above, secondaryveins 11–29 pairs with 2–8 mm spacing,55–75˚ from midrib, prominent or flat above,prominent beneath, clearly distinguishablefrom tertiary venation or not above, clearly
distinguishable beneath, straight, tertiary vena-tion prominent above or flat above and beneath,obscure or irregularly parallel to secondaryveins, intramarginal vein straight or onlyweakly looped, inset from margin.Inflorescence dichasial, 2.5–11.7 cm long withaxes 0.5–6.8 cm long and branches 0.7–0.8 mmwide, glabrous to densely puberulent; peduncle0.1–9.2 cm long, 0.7–0.8 mm wide, glabrous orpuberulent; pedicels 0.8–2.0 mm long,glabrous or densely puberulent, subtendingbracts persistent, bracts present on pedicel.Sepals 3.5–5.0 × 1.0–1.2 mm, 3.1–3.5 times aslong as wide, ovate or lanceolate, apex acumi-nate, ciliate or not, glabrous to densely puberu-lent outside, glabrous or puberulent on upperhalf inside. Corolla completely white; tube13–25 mm long, 1.5–2.3 mm wide, 1.7–1.9times as long as lobes, 4.3–6.8 times as long ascalyx, pubescent in upper part of tube above,around, and slightly below the stamens, throatglabrous, glabrous outside; lobes 7–15 ×1.8–6.7 mm, 2.2–4.7 times as long as wide,elliptic, apex acute to acuminate, not ciliate,glabrous outside and inside. Stamens inserted11.7–23.0 mm from corolla base, which is0.8–0.9 of corolla tube length in the rehydratedflowers measured; anthers 1.2–2.1 × 0.4–0.6
mm, 3.0–3.5 times as long as wide, apex0.6–1.8 mm from corolla throat; filaments0.4–0.8 mm long, pubescent. Disk 1 mm long,1.2–1.7 times as long as ovaries, glabrous, awl-shaped or lanceolate, apex acute or acuminate.Ovaries 0.6–0.8 mm high, sparsely pubescenton top; style 12.4 mm long; style head 0.6 mmlong. Fruit falcate with a small blunt hookedspur, 12–21 × 3.5–6.0 × 3.5–6.0 mm, spur 2mm long, sparsely puberulent.
Habitat: dry evergreen or deciduous forest at100–300 m altitude.
Additonal collections studied: LAOS:Vientiane Province: Houei Hin KannaWaterfall, 25 km N of Vientiane, Iwatsuki et al.IC 95-1594 (A); Nasaithong, Wongprasert etal. s.n. (BKF). THAILAND: Route 211 nearLoei border, Murata et al. T-50433 (BKF); PhuWua Wildlife Sanctuary, Pooma 1593 (A,BKF); Tham Phoon Falls, Wongprasert s.n.(BKF); Tadkham Falls, Wongprasert et al. s.n.(BKF).
This is a very distinctive species with smallwhite flowers, corolla lobes with acuminate toacute apices, and a congested inflorescencewith acuminate sepals and bracts.
FIGURE 2. Kopsia arborea Blume. A, habit; B, flower dissection; C, ovaries, disk, and base of style; D, fruit.From Hu 11985 (A, B, C) and Kostermans 7673 (D). Scale bars = 1 cm (A, B, D); 1 mm (C).
Kopsia pruniformis Reichb.f. & Zoll. exBakh.f., Blumea 6: 391. 1950; Sévenet etal., J. Ethnopharmacol. 41: 161. 1994.TYPE: INDONESIA. Java Timur,Rogodjampi, H. Zollinger 3832(Lectotype: NY, designated by Sévenet etal. [1994]; Isolectotypes: K, L, P).
Kopsia officinalis Tsiang & P.T.Li, ActaPhytotax. Sin. 11: 356. 1973; Sévenet etal., J. Ethnopharmacol. 41: 160. 1994.TYPE: CHINA. Yun-Ching-Hung,(=Junjinghong), Yunnan, Y.-M. Ting 54(Holotype: CANT; Photographs: A, P).
Tree to 14 m tall, to 30 cm dbh. Bark gray;inner bark pale brown. Branchlets glabrous orsparsely puberulent when young, sparselylenticellate or not. Leaves: petiole 3–10 mmlong, glabrous; blade 4.5–30.5 × 1.4–12.0 cm,1.9–5.7 times as long as wide, subcoriaceous tocoriaceous, elliptic, apex caudate to acuminatewith a blunt tip, base acute or cuneate, glabrousabove and beneath, midrib sunken to raised andwith a central groove above, secondary veins9–18 pairs with 4–13 mm spacing, 60–65˚ frommidrib, prominent above and beneath, clearlydistinguishable from tertiary venation aboveand beneath, straight or ascending near margin,tertiary venation prominent or flat above andbeneath, obscure or irregularly subperpendicu-lar to midrib and oblique to secondary veins,intramarginal vein strongly looped.Inflorescence dichasial, 4.8–15.3 cm long withaxes 2.0–12.5 cm long and branches 1.2–2.7mm wide, glabrous to sparsely puberulent;
peduncle 0.7–8.1 cm long, 2.1–2.7 mm wide,glabrous or puberulent in upper parts; pedicels0–5 mm long, glabrous to densely puberulent,subtending bracts persistent, bracts present onpedicel. Sepals 1.8–6.3 × 0.6–1.9 mm, 1.0–3.7times as long as wide, ovate, lanceolate oroblong, apex obtuse to acute, ciliate, glabrousto sparsely puberulent outside, glabrous orpuberulent on upper half inside. Corolla com-pletely white; tube 20.5–35.0 mm long, 1.6–2.2mm wide, 1.3–2.7 times as long as lobes,5.4–13.6 times as long as calyx, glabrous tosparsely pubescent around stamens and slightlybeneath inside, glabrous or sparsely puberulentat top of tube outside; lobes 7.0–21.5 × 3.4–6.5mm wide, 1.6–4.4 times as long as wide, ellip-tic or oblong, apex rounded to obtuse, ciliate orciliate only at lobe base, inside glabrous orsparsely pubescent in upper quarter, outsideglabrous. Stamens inserted 18.0–32.3 mm fromcorolla base which is 0.8–0.9 of corolla tubelength in the rehydrated flowers measured;anthers 1.2–1.7 × 0.5–0.8 mm, 2.0–3.4 times aslong as wide, apex 0.1–1.4 mm from corollathroat; filaments 0.8–1.2 mm long. Disk0.7–2.1 mm long, 1.1–1.9 times as long asovaries, glabrous, oblong, hourglass-shaped, orawl-shaped, apex shape variable and oftenquite complex, ranging from simply acuminateto rounded to horizontally V-shaped and flat ontop or horizontally V-shaped and retuse on top.Ovaries 0.9–1.2 mm high, glabrous to sparselypubescent all over; style 18–25 mm long; stylehead 0.8–1.1 mm long. Fruit oblique ellipsoidor subglobose, blue-black, 14.0–42.4 ×5.5–15.5 × 8–22 mm, only one carpel develop-ing (see note), spur absent, glabrous.
Habitat: grows in a wide range of foresttypes as an understory tree and at forest mar-gins on a wide range of soil types from sealevel to 1500 m altitude.
Geographical selection of the 185 collec-tions studied: ANDAMAN AND NICOBARISLANDS: South Andaman, Prain s.n. (A, M,US); Great Nicobar Island: Dhar Bay, Hore8750 (L). AUSTRALIA: Mossman Gorge, ca. 3miles SW of Mossman, Schodde 4170 (A, BRI,CANB, L). CHINA: Guangdong: Guangzhou,Yip 153 (BKF). Hainan: Yaichow, How 70531(A, K, NY, P, US). Hong Kong: Kadoorie Farm,Hu 11944 (A, K, PE). INDONESIA: Bali:
Brambang, Sarip 14 (L). Java: Java BaratProvince: Udjung Kulon Nature Reserve,Peutjang Island, Kostermans 47A (A, CANB,G, K, L, SAR). Kalimantan: Kalimantan TimurProvince, Balikpapan, Kostermans 7673 (A,BO, K, L). Lesser Sunda Islands: Flores,Manggarai, Golo Sengang, Schmutz 2750 (L).Sulawesi: Sulawesi Utara Province, SungaiIlanga, 220 km W of Manado, ca. 50 km inlandfrom Pangi, Burley, Tukirin et al. 3853 (A,BISH, E, K, KEP, L, MO). Sumatra: LampungProvince: Mt. Tanggamus, Jacobs 8043 (A,BISH, K, KEP, L, SING), Jacobs 8072 (A,BISH, K, L, SING). MALAYSIA: Penang:Waterfall Gardens, Haniff 3667 (BRI, SING,A). Sabah: Mt Kinabalu, Tenompok, Clemens& Clemens 26331 (A, BM, BO, L, K, NY, UC).Sarawak: Gunong Subis, Anderson S.16033 (K,L, SAN, SAR, SING). PHILIPPINES:Romblon, Elmer 12155 (A, BISH, BM, BO, E,G, L, MO, NY, US, Z). Mindoro Oriental:Subaan River, inland from San Teodoro,Ridsdale 1272 (A, BO, SAN). Negros Oriental:Lake Balinsasayao, Reynoso, Fuentes &Garcia 992 (GH). Leyte: Mt. Suiro, Sulit 21521(L, SING). THAILAND: Chiang MaiProvince: Ban Kong He, Kerr 3572 (ABD, BM,K). Nakhon Si Thammarat Province, KhaoLuang, Beusekom & Phengkhlai 901 (BKF, K,L, P). VIETNAM: Binh Tri Thien, Hue, MiiBach Ma Station, Poilane 31105 (P).
Typification: King and Gamble (1907) typi-fied Kopsia scortechinii with Scortechini 1878collected in Perak. Timmerman-Van derSleesen (1959) further lectotypified the namewith the SING duplicate. Sévenet et al. (1994)gave the following typification for this name:“Scortechini 57 b, 1878, holo-BM!, iso-SING!” It would appear that they thought thatthe two collections were duplicates and that1878 was a date, not a collecting number.However, it is clear from the style of the labelsand the known collecting dates of Scortechinithat 57b and 1878 are separate collections andthat only 1878 could be a type. In addition,King and Gamble list only 1878 in the proto-logue and 57 is listed as a syntype of K.larutensis, a specimen of which it undoubtedlyis. The type material of K. scortechinii fromCAL is clearly synonymous with K. arborea.Some further confusion has occurred throughMarkgraf’s (1973) distinction of K. scorte-chinii from K. arborea, partly based on the for-mer’s “Ovary with long white hairs,” even
though his second key suggests it has aglabrous ovary, which would agree with thespecimens and the original description.
In most species of Kopsia, the two carpelsboth develop into paired fruits, with the hooksfacing back toward each other. Sometimes onlyone of the carpels develops. In K. arborea,however, it would appear from the herbariumspecimens that only one of the two carpels everdevelops into a fruit. Although there is abun-dant herbarium material of this species, itshould be borne in mind that the fruits are oftenfallen ones placed in a packet, so this statementshould be treated with caution.
The collection Kostermans 7673 has some-how become confused, and specimens with thisnumber belong to both Alyxia reinwardtii andKopsia arborea.
The name Kopsia pitardii was applied byMerrill for Pitard’s concept of K. cochinchinen-sis Kuntze, the type of which is a specimen ofTabernaemontana divaricata. Although Merrillreferred to it as a nom. nov., it is actually a newspecies by Merrill, validated by reference backto Pitard’s description of K. cochinchinensis.Sévenet et al. (1994) have effectively lecto-typified the Paris element of the collectionPierre 32. In Paris there are three specimens ofPierre 32, but only one of them is labelled as K. cochinchinensis, the name taken up byPitard (1933), on which K. pitardii was based.Therefore, it is unnecessary to make a secondstep lectotypification for this particular specimen.
Tree to 10 m tall, to 46 cm dbh. Bark gray oryellow, smooth; inner bark white. Branchletsglabrous, sparsely lenticellate, weakly winged.Leaves: petiole 5–7 mm long, glabrous; blade6.5–24.5 × 2.0–10.2 cm, 2.1–3.3 times as longas wide, subcoriaceous to coriaceous, mostlydrying reddish-brown, elliptic, apex longacuminate with a blunt tip, base acute tocuneate, glabrous above and beneath, midribshallowly sunken or raised and with a centralgroove above, secondary veins 9–16 pairs with
100 HARVARD PAPERS IN BOTANY Vol. 9, No. 1
6–18 mm spacing, 65–70˚ from midrib, promi-nent to sunken above and prominent beneath,clearly distinguishable from tertiary venationabove and beneath, straight, ascending nearmargin or curved ascending from midrib, ter-tiary venation prominent or flat above andbeneath, obscure or irregularly subperpendicu-lar to midrib and oblique to secondary veins,intramarginal vein straight, weakly or stronglylooped, inset from margin. Inflorescence withdichasial branching followed by cincinnatebranches, 10.7–20.0 cm long with axes 7–16cm long and branches 2.0–2.3 mm wide,densely puberulent; peduncle 2.7–9.3 cm long,1.9–2.3 mm wide, puberulent; pedicels ca. 1.2mm long, densely puberulent, subtendingbracts persistent. Sepals 2.3–3.7 × 1.4–2.2 mm,1.60–1.75 times as long as wide, oblong, apexrounded to obtuse, ciliate, densely puberulentoutside, glabrous inside. Corolla completelywhite; tube 22–35 mm long, 1.4 mm wide,1.1–1.4 times as long as lobes, 5.9–13.5 timesas long as calyx, pubescent around and beneathstamens and in throat, glabrous or sparselypuberulent at top of tube outside; lobes 16–28× 6.2–7.0 mm, 2.6–4.0 times as long as wide,elliptic, apex rounded, ciliate only at lobe base,glabrous outside and inside. Stamens inserted19–21 mm from corolla base, which is ca. 0.7
of corolla tube length in the rehydrated flowersmeasured; anthers 1.9–2.6 × 0.5–0.6 mm wide,3.8–4.6 times as long as wide, apex 5.2–5.8mm from corolla throat; filaments 0.8–0.9 mmlong, pubescent. Disk 1.1–1.5 mm long,0.9–1.7 times as long as ovaries, glabrous, awl-shaped, narrowly deltoid or lanceolate, apexacuminate. Ovaries 0.9–1.5 mm high, denselypubescent on top or densely pubescent all over;style 17–20 mm long; style head 1.1–1.4 mmlong. Fruit sparsely puberulent, with a blunthooked spur, 12–17 × 5.4–6.0 × 7–10 mm, spur3.5–6.0 mm long.
Distribution: Borneo (Sabah).Habitat: in a variety of evergreen forest types,
often on sandy soils from 0–900 m altitude.Selection of the 83 collections studied:
MALASIA. Sabah, Lahad Datu District, UluSegama, Argent et al. 108232 (E, K, KEP,L, SAN, SAR); Danum Valley, MadaniSAN116456 (K, L, SAN).
Possibly related to Kopsia pauciflora but dif-fering from that species in the stamens beingsituated somewhat lower in the corolla tube andin the generally much more densely pubescentand often much longer inflorescences. In someareas in Sabah, such as in Danum Valley, itwould appear to be the most common species.
FIGURE 3. Kopsia dasyrachis Ridl. A, habit; B, flower dissection; C, ovaries, disk, and base of style; D, fruit.From Gibet 37111 (A, B, C) and Enggoh 10435 (D). Scale bars = 1 cm (A, B, D); 1 mm (C).
4. Kopsia deverrei L.Allorge, Phytologia 59:93. 1986; Sévenet et al., J. Ethnopharmacol. 41:151. 1994; I.M.Turner, Gard. Bull. Sing. 47:127. 1995. TYPE: MALAYSIA. Johor, 87 kmmilestone from Mersing to Johor Bahru,Deverre 25 (Holotype: P; Isotype: P). Map 4.
Tree to 10 m tall. Branchlets glabrous,sparsely lenticellate or not, terete or weaklyangled. Leaves: petiole 6–17 mm long,glabrous; blade 4.6–14.5 × 1.1–5.5 cm, 2.1–3.8times as long as wide, papery or subcoriaceous,elliptic, apex caudate or short to long acumi-nate with a blunt tip, base acute or cuneate,glabrous above and beneath, midrib shallowlysunken above, secondary veins 7–11 pairs with3–15 mm spacing, 45–60˚ from midrib, promi-nent to sunken above, prominent beneath,clearly distinguishable from tertiary venationabove and beneath, ascending near margin orcurved ascending from midrib, tertiary vena-tion prominent above, prominent or flatbeneath, subperpendicular to midrib andoblique to secondary veins and also somewhatreticulate, intramarginal vein straight tostrongly looped, inset from margin.Inflorescence dichasial, 4.5–4.8 cm long withaxes 0.9–2.4 cm long and branches 0.8–0.9 mmwide, glabrous; peduncle 0.2–2.6 cm long,
0.9–1.4 mm wide, glabrous; pedicels 1.0–3.5mm long, glabrous or sparsely puberulent; sub-tending bracts persistent; bracts present onpedicel. Sepals 1.6–1.7 × 1.4 mm, 1.1–1.2times as long as wide, ovate, apex rounded toobtuse, ciliate, glabrous outside and inside.Corolla white with a red “eye”; tube 27–37 mmlong, 2 mm wide, 1.5–2.1 times as long aslobes, 15.9–23.1 times as long as calyx, pubes-cent in upper part of tube above, around, andslightly below the stamens, throat pubescent,glabrous outside; lobes 13–24 × 4.5–4.8 mm,2.7–5.3 times as long as wide, elliptic oroblong, apex rounded, ciliate, pubescent at verybase of lobes, glabrous outside. Stamensinserted 22–32 mm from corolla base, which isca. 0.9 of corolla tube length in the rehydratedflowers measured; anthers 1.3–1.5 × 0.6 mm,2.2–2.5 times as long as wide, 0.8–1.4 mmfrom corolla throat; filaments ca. 0.8 mm long,pubescent. Disk 1.0–1.1 mm long, 1.1–1.2times as long as ovaries, glabrous, awl-shaped,apex acute or acuminate. Ovaries ca. 0.9 mmhigh, densely pubescent on top; style 21.5–31.0mm long; style head 0.8–0.9 mm long. Fruitunknown.
Distribution: Peninsular Malaysia (Johor).Habitat: from 150 to 305 m altitude.
In this species, the corolla tube is particularlydensely pubescent behind the anthers.Specimens of this species were included inKopsia singapurensis by Timmerman-Van derSleesen (1959) and by Markgraf (1973),although Timmerman-Van der Sleesenexpressed her doubt that they belonged there.This species is close to K. singapurensis, dif-fering from it in the remarkably long petioles ascompared with the length of the leaf blade, themore delicate inflorescences, and the shorter,more obtuse sepals. However, there are alsosome specimens of K. singapurensis, mostnotably Hamid 10875 (KEP), with the shorterpetioles, large leaves, and venation patterns ofK. singapurensis but with the smaller flowersapproaching those of K. deverrei, which some-what bridge the gap between them. I havemaintained K. deverrei here, as the material ofthis species is readily recognized by the longpetioles and different leaf size and shape.However, more-detailed population studies ofthe two species may lead to a reevaluation ofthe distinctions between them.
5. Kopsia flavida Blume, Rumphia 4: 28. 1849;Merr., Phil. J. Sc. 29: 412. 1926; Miq., Fl. Ind.Bat. 2: 410. 1857; Corner, Wayside TreesMalaya 1: 145. 1952; Sleesen, Fl. Mal. Misc.Rec. 1: 10. 1959; Markgr., Blumea 20: 423.1973; Sévenet et al., J. Ethnopharmacol. 41:151. 1994. TYPE: NEW GUINEA. Sine loc.[but likely West Papua], Zippelius s.n.(Lectotype: L [898.110–336], designated byTimmerman-Van der Sleesen [1959];Isolectotypes: L [898.110–334, 898.110–335]).Map 5; Fig. 4.Synonyms: Calpicarpum albiflorum Teijsm. &
Binn., Tijdschr. Nederl. Ind. 25: 402.1863. TYPE: INDONESIA. Seram, J.E.Teijsmann HB5035 (Lectotype: BO, des-ignated by Sévenet et al. [1994]; Isotype:L). Kopsia albiflora (Teijsm. & Binn.)Boerl., Handl. Fl. Ned. Ind. 2: 395. 1899;Sévenet et al., J. Ethnopharmacol. 41:149. 1994; Kopsia fruticosa var. albiflora(Teijsm. & Binn.) King & Gamble, J. As.Soc. Beng. 4(2): 431. 1907.
Calpicarpum ornatum W.Bull, RetailList 199: 12. 1884; Fraser & Hemsl.,Johns Gard. Dict. new ed. 156. 1917;Mabberley, Taxon 34: 456. 1985. Noillustration or specimen has been found
FIGURE 4. Kopsia flavida Blume. A, habit; B, flower dissection; C, ovaries and disk; D, fruit. From Brass 3110(A) and Takeuchi 9028 (B, C, D). Scale bars = 1 cm (A, B, D); 1 mm (C).
that could be type material. However,from the description and its provenance itis clearly Kopsia flavida.
Kentrochrosia monocarpa Laut. &K.Schum., Fl. Schutzgeb. Südsee 506.1900; Markgr., Bot. Jahrb. 61: 195. 1927;Markgr., Nov. Guinea 16(2): 283. 1927;Merr. & L.M.Perry, Phil. J. Sci. 76: 20.1941. TYPE: PAPUA NEW GUINEA.Oertzen Mountains, C.A.G. Lauterbach2180 (Holotype: B, not found, presum-ably destroyed; Schumann & Lauterbach,Fl. Schutzgeb. Südsee t. XVIII, 1900,Lectoype, designated here). The originaltype specimen could not be found, but itis clear from the collecting locality, thedescription, and the illustration that itbelongs to Kopsia flavida.
Kopsia grandiflora Merr., Phil. J. Sci. 20:435. 1922; Sévenet et al., J.Ethnopharmacol. 41: 154. 1994. TYPE:PHILIPPINES. Luzon, Camarines SurProvince, Paracale, M. Ramos, G.E.Edaño 33691 (Lectotype: US, designatedhere; Isolectotypes: A, K, P).
Kopsia triangularis Quisumb. & Merr.,Phil. J. Sci. 37: 191. 1928; Sévenet et al.,J. Ethnopharmacol. 41: 165. 1994.TYPE: PHILIPPINES. Mindanao,Surigao, C.A. Wenzel 2648 (Holotype:UC; Isotypes: A, BO, BR, G, M, MO, Z).Kentrochrosia triangularis (Quisumb. &Merr.) Merr. & L.M.Perry, Phil. J. Sci.76: 21. 1941.
Tree to 9.15 m tall, to 27.5 cm dbh. Barkashy-brown, brown, or gray, smooth; inner barkstraw colored. Branchlets glabrous, sparselylenticellate or not. Leaves: petiole 3–10 mmlong, glabrous; blade 4.5–25.0 × 1.9–8.9 cm,1.7–4.9 times as long as wide, subcoriaceous tocoriaceous, elliptic, apex acuminate with ablunt tip, base acute or cuneate, glabrous aboveand beneath, midrib deeply or shallowly
sunken above, secondary veins 8–24 pairs with3–15 mm spacing, 60–75˚ from midrib, promi-nent above and beneath, clearly distinguishablefrom tertiary venation above and beneath,straight or ascending near margin, tertiaryvenation prominent above and beneath, reticu-late, intramarginal vein obscure or more or lessstraight and inset from margin. Inflorescencedichasial or cincinnate, 6.0–6.5 cm long withaxes 1.0–3.5 cm long and branches 1.2–2.2 mmwide, glabrous or puberulent in upper parts;peduncle 0.2–1.0 cm long, 1.2–2.2 mm wide,glabrous; pedicels 1.5–3.5 mm long, glabrousor sparsely puberulent, subtending bracts per-sistent, bracts present on pedicel. Sepals1.6–3.0 × 0.9–2.4 mm, 1.1–2.2 times as long aswide, ovate or oblong, apex rounded to acute,ciliate, glabrous outside and inside. Corollawhite with a red “eye” or pale pink with adarker pink “eye”; tube 26–38(– 49) mm long,2.2–3.0 mm wide, 1.2–1.8 times as long aslobes, 11.7–16.3 times as long as calyx, pubes-cent around and beneath stamens and in throat,rarely glabrous, glabrous outside; lobes 16–31× 6.1–12.5 mm, 1.3–3.5 times as long as wide,elliptic or obovate, apex rounded to obtuse, cil-iate or not, glabrous outside and inside.Stamens inserted 23–40 mm from corolla base,which is 0.8–0.9 of corolla tube length in therehydrated flowers measured; anthers 1.5–2.6 ×0.5–0.8 mm, 3.0–3.7 times as long as wide,apex 0.9–2.5 mm from corolla throat; filaments0.7–1.1 mm long, pubescent. Disk 1.2–1.9 mmlong, 0.9–1.4 times as long as ovaries,glabrous, oblong, awl-shaped, narrowly deltoidor lanceolate, apex acute to acuminate. Ovaries1.2–1.5 mm high, glabrous or, rarely, verysparsely pubescent on top; style 23–38 mmlong; style head 1.0–1.4 mm long. Fruit with asharp deltoid spur, 27–35 × 10–14 × 3.0–5.5mm, spur 8–13 mm long, glabrous.
Habitat: in a wide range of forest types fromswamp forest to lowland evergreen forest, onlimestone, ultrabasic, and poorly to well-drained soils from 0 to 270 m altitude.
Geographical selection of the 86 collec-tions studied: CAROLINE ISLANDS: Palau,Babeldaob, Aimiyou, Kanehira 502 (FU, NY).INDONESIA: West Papua: Merau River, Southof Senajo, Royen 4674 (L). Moluccas: Buru,Vriese s.n. (L); Wae Toni, Eyma 3171 (A,SING, K, L); Kai Is, Jaheri 8 (BO, L); Seram,Vriese, Teijsmann s.n. (L). PAPUA NEW
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GUINEA: Western Province: Wuroi, Brass5789 (A, BM, BO, BRI); West New BritainProvince: Gasmata, Kerenga, Obedi LAE62338(L); Manus: Lorengau, Kerenga et al.LAE77481 (CANB, L). PHILIPPINES: ManiKani Island, Quisumbing 2049 (A). Surigao:Lake Mainit, Ramos & Convocar 83386 (A).Eastern Samar: Balangiga, Madulid et al. 1222(L). SOLOMON ISLANDS: North Solomons:San Jorge, Talise Village, Hunt RSS2730 (A, K,L). Santa Cruz Islands: Vanikoro Island,Kajewski 581 (A, BRI, K, US). San Cristobal:Star Harbour, Brass 3110 (A, BO, BRI, L).Santa Isabel: Allardyce Harbour, Kinifu, SusuiBSIP8279 (K, L, SING). VANUATU: Tisbel,Hallé RSNH 6374 (K, L, P). Efate, Burton 58(A).
Rutten 1895 from Seram has a corolla tubevery much longer than all other known speci-mens of this species. Unfortunately there are nolabel notes on corolla color. In other charactersit is similar to the other specimens from Seram.Additional collections may lead to a reappraisalof this decision.
MAP 6. Distribution of Kopsia fruticosa (Roxb.) A.DC.
Tree to 6 m tall. Bark mostly grayish.Branchlets sparsely to densely puberulent,glabrescent when older, not lenticellate, weaklyangled. Leaves: petiole 5–12 mm long,glabrous; blade 5.4–22.0 × 2.5–10.2 cm,1.7–3.5 times as long as wide, subcoriaceous orcoriaceous, elliptic, apex short to long acumi-nate with blunt tip, base obtuse to cuneate,glabrous above and beneath, midrib deeplysunken to flat above, secondary veins 8–17pairs with 4–21 mm spacing, 50–70˚ frommidrib, slightly sunken to prominent above,prominent beneath, clearly distinguishablefrom tertiary venation above and beneath,curved ascending from midrib, tertiary vena-tion prominent or flat above and beneath, sub-perpendicular to midrib and oblique tosecondary veins or obliquely scalariform, usu-ally also somewhat reticulate, intramarginalvein obscure or straight to strongly looped andinset from margin. Inflorescence dichasial,4.9–9.5 cm long with axes 1.1–5.5 cm long andbranches 1.1–1.9 mm wide, sparsely to denselypuberulent; peduncle 0.3–4.0 cm long, 1.9–2.9mm wide, puberulent at least in upper parts;pedicels 1.4–5.0 mm long, sparsely to denselypuberulent; subtending bracts persistent; bractspresent on pedicel. Sepals 1.7–2.5 × 1.2–2.2mm, 1.0–1.7 times as long as wide, ovate oroblong, apex emarginate to obtuse, ciliate,glabrous to densely puberulent, glabrous orpuberulent on upper half inside. Corolla mostlywhite or pale pink at the base of the tube andbecoming pinker higher up, lobes whitish-pinkto pink with a darker pink to almost red throat,more rarely whole corolla only pale pinkish;tube 25–37 mm long, 1.8–3.2 mm wide,1.5–2.5 times as long as lobes, 11.6–21.2 timesas long as calyx, pubescent around stamens andslightly beneath inside, throat pubescent,glabrous outside; lobes 10–33 × 5.5–16.0 mm,1.6–2.4 times as long as wide, elliptic, apexrounded, ciliate or ciliate only at lobe base,glabrous outside and inside. Stamens inserted20.0–33.5 mm from corolla base, which is0.8–0.9 of corolla tube length in the rehydratedflowers measured; anthers 1.9–2.5 × 0.5–0.8mm, 2.9–3.8 times as long as wide, 1.4–2.4 mmfrom corolla throat; filaments 0.6–0.9 mm long,pubescent. Disk 1.0–1.7 mm long, 0.7–1.3times as long as ovaries, glabrous, oblong orawl-shaped, apex acute, acuminate or irregu-larly toothed and acuminate. Ovaries 1.2–1.6mm high, densely pubescent; style 20.5–32.0
mm long; style head 1.1–1.5 mm long. Fruitfalcate with a blunt hooked spur, ca. 17 × 5 × 7mm, spur ca. 3 mm long, densely puberulent.
Distribution: Burma (Tenasserim), alsowidely cultivated. The original description sug-gests the plant comes from Pegu, but no collec-tions have been seen from that far north.
Habitat: no habitat information has beenprovided on the collections made in the wild.
Typification: The basionym of this species,Cerbera fruticosa, was originally published inRoxburgh (1814), but without a description,and later described in Ker Gawler’s BotanicalRegister (1819). Roxburgh (1814) and KerGawler (1819) note that it originally came fromPegu, Burma, a site much farther north than anyof the known collection localities. Ker Gawlerfurther explains that it was introduced as a hor-ticultural shrub to the Calcutta Botanic Gardenand eventually to the United Kingdom. KerGawler gives credit to Roxburgh for the nameand the description; the validating descriptionitself is followed by “Roxb. MSS” and thewording is the same as that in Roxburgh’s FloraIndica (1824). Curiously, Roxburgh has notbeen credited for the name in the bulk of the lit-erature, with the authority invariably given asKer Gawler (or variations on his name). Itappears to me, however, that Ker Gawler hashimself acknowledged that this species wasnamed by Roxburgh and that the validatingdescription comes from a Roxburgh manu-script, and therefore the authority should ratherbe Cerbera fruticosa Roxb. Roxburgh creditsthis name to himself but with Ker Gawler(1819) as the place of publication in hisaccount of Cerbera fruticosa in Flora Indica(1824). This raises the issue of typification.Sévenet et al. (1994) typified this species withthe illustration of a cultivated plant growing inFulham in Southeast England in Ker Gawler’s1819 account. However, given that Roxburghbased his name on material cultivated in theCalcutta Botanic Garden, the material fromCalcutta must be the type. Forman (1997) didnot mention this species in his list of Roxburghspecies, and I have been unable to find any
108 HARVARD PAPERS IN BOTANY Vol. 9, No. 1
herbarium material that could unequivocally beconsidered original (although there is a speci-men in BM labelled as Cerbera fruticosa inRoxburgh’s handwriting but with no otherinformation on the sheet). Sealy (1956) andSanjappa et al. (1994) note that illustrationsexist in the Roxburgh Flora Indica collectionsat Kew and Calcutta. Forman (1997) makesclear that these illustrations were made undersupervision by Roxburgh and that there was avery close link between them and the descrip-tions in Flora Indica. Hence these illustrationscan be considered part of the original material,and a lectotype can be made of the Kew illus-tration, number 2200. The illustration has alsobeen reproduced by Wight (1841) as t. 431under the name Calpicarpum roxburghii.
The large majority of specimens known ofthis species are from cultivation throughout thetropics. Only the specimens that are likely tohave come from the wild have been mapped.Only one fruiting specimen has been seen,which is quite curious for such a commonlycollected plant, albeit mostly cultivated.
There are a number of specimens labelledKopsia vincaeflora in Blume’s handwriting inthe Leiden herbarium. It is not clear whetherthese are duplicates of a single collection or if
they represent multiple collections, and there-fore no isolectotypes can be distinguished fromthe syntypes with any degree of certainty.
7. Kopsia grandifolia D.J.Middleton, sp. nov.TYPE: MALAYSIA. Johor, Sungai Mupar, off32nd mile Johor Bahru–Mersing Road, H.M.Burkill 4230 (Holotype: A; Isotypes: K, L,SAR, SING). Map 7; Fig. 5.
Frutex 2–3 m altus, ramulis puberulis. Foliaelliptica, 19.0–29.5 × 6.1–12.1 cm, nervis24–32 paribus. Corollae tubus 15.5–18.0 mm,lobis 10–12 × 1.6–2.3 mm.
Tree to 3 m tall. Branchlets sparsely puberu-lent, sparsely lenticellate or not, terete orweakly angled. Leaves: petiole 7–11 mm long,sparsely to densely pubescent; blade 19.0–29.5× 6.1–12.1 cm, 2.4–5.2 times as long as wide,papery or subcoriaceous, broadly elliptic orelliptic, apex caudate, rarely to short acuminatewith a blunt tip, base obtuse to cuneate,glabrous or puberulent on midrib above,puberulent on midrib and sometimes on majorveins beneath, at least when young, midribshallowly sunken to flat above, secondary veins24–46 pairs with 3–15 mm spacing, 65–80˚from midrib, prominent or flat above, prominentbeneath, clearly distinguishable from tertiary
MAP 7. Distribution of Kopsia grandifolia D.J.Middleton (•) and Kopsia lapidilecta Sleesen (★).
110 HARVARD PAPERS IN BOTANY Vol. 9, No. 1
FIGURE 5. Kopsia grandifolia D.J.Middleton. A, habit; B, flower dissection; C, ovaries, disk, and base of style;D, fruit. From Burkill HMB.4230 (A, B, C) and David 265 (D). Scale bars = 1 cm (A, B, D); 1 mm (C).
venation above and beneath, straight or ascend-ing near margin, tertiary venation prominent or flat above, prominent beneath, parallel tosecondary veins or subperpendicular to midriband oblique to secondary veins, usually alsosomewhat reticulate, intramarginal veinstraight or looped, inset from margin.Inflorescence dichasial, with or without longerunbranched branches, 3.2–8.0 cm long withaxes 1.7–6.5 cm long and branches 1.5–1.9 mmwide, sparsely to densely puberulent; peduncle0.7–1.0 cm long, puberulent, pedicels ca. 3 mmlong, densely puberulent; subtending bractspersistent; bracts present on pedicel. Sepals 1.6× 1.5–2.2 mm, 0.7–1.1 times as long as wide,ovate, apex rounded, ciliate, densely puberu-lent, glabrous inside. Corolla buds purple, yel-low when open; tube 15.5–18.0 mm long, ca.1.8 times as long as lobes, ca. 9.7 times as longas calyx, densely pubescent only beneath sta-mens and becoming less so further down inupper half, throat glabrous, glabrous outside;lobes 10–12 × 1.6–2.3 mm, 4.9–6.7 times aslong as wide, linear to oblong, apex rounded toacute, ciliate or not, glabrous outside andinside. Stamens inserted at ca. 15 mm fromcorolla base, which is ca. 0.8 of corolla tubelength in the rehydrated flower measured,anthers 2.6–2.7 × 0.6 mm, 4.3–4.5 times aslong as wide, apex 0.4–1.1 mm from corollathroat. Disk 1.0–1.1 mm long, ca. 0.8 times aslong as ovaries, glabrous, awl-shaped or del-toid, apex acuminate. Ovaries 0.8–0.9 mmhigh, glabrous; style 1.5–15.0 mm long (seenote); style head 0.9–1.1 mm long. Fruit falcatewith a blunt hooked spur in the lower part ofthe fruit, 18.5–19.0 × 9.5–10 × 4 mm, hook 2–3mm long, sparsely puberulent.
Distribution: Peninsular Malaysia and proba-bly the Anambas Islands (see comment below).
Habitat: in forest to 60 m altitude.Additional collections studied: INDONE-
SIA: Anambas Islands, North Jemaja, GunongDatoh, Henderson 20395 (SING). MALAYSIA:Johor: Mersing, David 265 (P), Teo et al. 1199(P); Sungai Mupah, 53 km from Johor Barutowards Mersing, Teo & Pachiapu 1127 (K, L);Kluang Forest Reserve, Hutan Simpan Gunong
Belumut, Teo & Pachiapu 1143 (K, L, SING).This new species is close to Kopsia laruten-
sis, differing from it in its much larger flowers,leaves generally pubescent on the lower surface(at least when young), glabrous ovaries, andlarger and longer fruits. Both species have veryrestricted distributions: K. larutensis in theMalaysian state of Perak and K. grandifolia inthe Malaysian state of Johor and probably thenearby Indonesian Anambas Islands. Henderson20395 from the Anambas Islands is a paratypeof K. lapidilecta, the type of which is from theNatuna Islands. It is a rather poor specimenwithout a corolla. However, it does have leaveslike K. grandifolia, pubescent when young, theremnants of the inflorescence are pubescent, anda comment on the label notes that the now-miss-ing corolla lobes are narrow. Kopsia lapidilectaalso has rather narrow corolla lobes but hasglabrous leaves and a glabrous inflorescence.
David 265 has an extremely short style,shorter than any other observed in the genus.The type collection of the species, BurkillHMB4230, has a style that places the style headat the same level as the stamens, as is the casein all the other species. Because there are sofew specimens of this species, I am unsurewhether the short style of David 265 is just anaberrant character or whether it has some polli-nation function.
8. Kopsia griffithii King & Gamble, J. As. Soc.Beng. 74(2): 432. 1907; Ridl., Fl. Mal. Pen. 2:337. 1923; Sleesen, Fl. Mal. Misc. Rec. 1: 12.1959; Whitmore, Tree Fl. Mal. 2: 19. 1972;Markgr., Blumea 20: 421. 1973; Sévenet et al.,J. Ethnopharmacol. 41: 154. 1994; I.M.Turner,Gard. Bull. Sing. 47: 127. 1995. TYPE:MALAYSIA. Malacca sine loc., W. Griffith s.n.(Lectotype: CAL, designated by Sévenet et al.[1994]; Isolectotypes BO, K, L, M). Map 8;Fig. 6.Synonym: Kopsia griffithii var. paucinervia
King & Gamble, J. As. Soc. Beng. 74(2):432. 1907. TYPE: MALAYSIA. Peraksine loc., King’s Collector 10707(Lectotype: UC, designated here;Isolectotype: BO).
1a. Pedicels glabrous; corolla tube > 30 mm long, lobes > 19 mm long . . . . . . . . . . . . . . . . . . . . . . . . . var. griffithii1b. Pedicels densely pubescent; corolla tube < 25 mm long, lobes < 15 mm long . . . . . . . . . . . . . . . . var. pubescens
Kopsia griffithii King & Gamble var. griffithiiTree to 5 m tall; to 24 cm dbh. Bark ashy-
brown or gray. Branchlets glabrous, not lenti-cellate or sparsely lenticellate. Leaves: petiole3–10 mm long, glabrous; blade 5.7–20.0 ×1.6–8.0 cm, 2.0–5.3 times as long as wide,papery to subcoriaceous, elliptic to oblong,apex caudate or long acuminate with a blunttip, base acute or cuneate, glabrous above andbeneath, midrib raised and with a centralgroove to sunken above, secondary veins 9–28pairs with 1–10 mm spacing, 45–80˚ frommidrib, clearly distinguishable or not from ter-tiary venation above and beneath, prominentbeneath, prominent or flat above, straight orascending near margin, tertiary venation promi-nent above and beneath, reticulate or subper-pendicular to midrib and oblique to secondaryveins, intramarginal vein straight or onlyweakly looped, inset from or right at margin.Inflorescence dichasial, lax, 5–12 cm long ofwhich the axes are 1.2–7.0 cm long andbranches 0.7–1.0 mm wide, glabrous; peduncle0.5–6.0 cm long, 0.9–1.5 mm wide, glabrous;pedicels 1.0–3.5 mm long, glabrous, subtend-ing bracts persistent, bracts present or absent onpedicel. Sepals 1.4–1.8 × 0.7–1.4 mm, 1.1–2.1times as long as wide, ovate or oblong, apexrounded to acute, ciliate, glabrous outside andinside. Corolla completely white or white with
yellow “eye”; tube 31–40 mm long, 1.6–2.4mm wide, 1.6–2.2 times as long as lobes,19.4–26.0 times as long as calyx, pubescentaround and beneath stamens and in throat,glabrous outside; lobes 16–23 × 4.5–7.0 mm,2.5–4.6 times as long as wide, elliptic, apexobtuse or acute, ciliate or not, glabrous outsideand inside. Stamens inserted 26.5–35.0 mmfrom corolla base, which is 0.8–0.9 of corollatube length in the rehydrated flowers measured;anthers 2.0–2.3 × 0.5–0.7 mm, 3.3–4.6 times aslong as wide, apex 1.7–2.6 mm from corollathroat; filaments 0.7–1.0 mm long, pubescent.Disk 0.8–1.4 mm long, 0.8–1.2 times as long asovaries, pubescent, awl-shaped, apex acute.Ovaries 0.8–1.4 mm high, densely pubescenton top or densely pubescent all over; style26–32 mm long; style head 0.8–1.2 mm long.Fruit falcate with a small spur, 19–21 × 7–8 × 3mm, spur 3–4 mm long; sparsely puberulent.
Distribution: Peninsular Malaysia.Habitat: in primary or secondary forest to
800 m altitude.Selection of the 35 collections studied:
MALAYSIA: Selangor: Templer Park, Stone5956 (MO, US), David 268 (P), Teo &Pachiapu 298 (CANB, K, L, SING), 1130 (K,L, SING); Ipoh Highway, 15 km NW of KualaLumpur, Worthington 13072 (L, MO, NY).
The only material I was able to trace of
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MAP 8. Distribution of Kopsia griffithii King & Gamble.Var. griffithii (•), and var. pubescens D.J.Middleton (★).
FIGURE 6. Kopsia griffithii King & Gamble. A, habit; B, flower dissection; C, ovaries, disk ,and base of style;D, fruit. From Kasim 590 (A) and Wyatt-Smith 93110 (B, C, D). Scale bars = 1 cm (A, B, D); 1 mm (C).
King’s Collector 10707, the type of Kopsiagriffithii var. paucinervia King & Gamble, wasfrom BO and UC. I have seen no material fromCAL or K, where one would expect the typematerial to be. Therefore, the name has beenlectotypified by the UC specimen.
Kopsia griffithii King & Gamble var. pubes-cens D.J.Middleton, var. nov. TYPE:MALAYSIA. Johor, Sungai Hula Sembrong,30 October 1892, Lake & Kelsall s.n.(Holotype: SING).
A Kopsia griffithii var. griffithii pedicellispubescentibus corollis parvioribus differt.
Height unknown. Branchlets glabrous, notlenticellate, weakly angled. Leaves: petiole 5–7mm long, glabrous; blade 11.3–17.0 × 3.9–7.2cm, 2.6–2.8 times as long as wide, coriaceous,elliptic, apex long acuminate with a blunt tip,base acute, glabrous above and beneath, midribshallowly sunken above, secondary veins18–20 pairs with 5–14 mm spacing, 60–70˚from midrib, ascending near margin, prominentand not clearly distinguishable from tertiaryvenation above, tertiary venation prominentand reticulate or subperpendicular to midriband oblique to secondary veins above, intra-marginal vein straight or only weakly looped,inset from margin. Inflorescence dichasial, ca.11 cm long with axes ca. 9.3 cm long andbranches ca. 1 mm wide, densely puberulent;peduncle ca. 5 cm long, ca. 2.5 mm wide,puberulent in upper parts; pedicels 2.1–3.5 mmlong, densely puberulent; subtending bractspersistent; bracts absent on pedicel. Sepals ca.2.0 × 1.2 mm, ca. 1.7 times as long as wide,ovate, apex acute, ciliate, sparsely puberulentoutside, glabrous inside. Corolla tube 18–20mm long, ca. 1.6 mm wide, ca. 1.9 times aslong as lobes, ca. 9 times as long as calyx,pubescent around stamens and slightly beneathinside, throat pubescent, glabrous outside;lobes ca. 9.5 × 2.5 mm, ca. 3.8 times as long aswide, elliptic, apex obtuse, not ciliate, glabrousoutside and inside. Stamens inserted at ca. 18mm from corolla base which is ca. 0.9 ofcorolla tube length in the rehydrated flowermeasured; anthers ca. 1.4 × 0.4 mm, ca. 3.5times as long as wide, apex ca. 0.6 mm fromcorolla throat. Disk ca. 0.6 mm long, ca. 0.7times as long as ovaries, pubescent, awl-shaped, apex acute. Ovaries ca. 0.9 mm high,densely pubescent all over; style ca. 16.5 mmlong; style head ca. 8 mm long. Fruit unknown.
Distribution: Peninsular Malaysia (Johor).
Habitat: not recorded.The single specimen known of this taxon is
an old Lake & Kelsall s.n. specimen in SINGcollected on 30 October 1892. It is unfortu-nately not a very good specimen, and I havecreated this variety somewhat reluctantly. It is,however, quite different from typical Kopsiagriffithii, and future collections of var. pubes-cens may determine that it deserves specificstatus. The main differences from var. griffithiiare in the rather more lax and many-floweredinflorescence, the densely pubescent pedicels,and the smaller flowers. It shares with var. grif-fithii the lax inflorescence and a pubescentdisk, and the two are the only taxa in the genusthat have this latter character.
9. Kopsia hainanensis Tsiang, Sunyatsenia 2:111. 1934; Sévenet et al., J. Ethnopharmacol.41: 154. 1994. TYPE: CHINA. Hainan:Yaichow, F.C. How 70532 (Lectotype: IBSC,designated here; Isolectotypes: A, CANT, E, K,P). Syntype: N.K. Chun & C.L. Tso 43853 (A,NY). Map 9.
Tree to 2 m tall. Bark gray. Branchletsglabrous, sparsely lenticellate or not. Leaves:petiole 4–8 mm long, glabrous; blade 3.0–15.3× 1–4 cm, 2.2–5.8 times as long as wide,papery or subcoriaceous, elliptic, apex acumi-nate with a blunt tip, base cuneate, glabrousabove and beneath, midrib raised and with acentral groove, flattened or shallowly sunkenabove, secondary veins 16–22 pairs with 2–10mm spacing, 55–75˚ from midrib, prominentabove, prominent or not beneath, clearly distin-guishable or not from tertiary venation aboveand beneath, tertiary venation prominent above,prominent or not beneath, reticulate, intramar-ginal vein straight or only weakly looped,slightly inset or right at margin. Inflorescencedichasial, 4.0–4.5 cm long, of which the axesare 0.4–1.0 cm long and branches 1.1–1.4 mmwide, glabrous; peduncle 0.1–0.3 cm long,1.1–1.4 mm wide, glabrous; pedicels 1.7–4.0mm long, glabrous, bracts present on pedicel.Sepals 1.2–1.3 × 0.7–1.1 mm, 1.1–1.7 times aslong as wide, ovate, apex obtuse to acute, cili-ate or not, glabrous outside, glabrous orpuberulent at tips inside. Corolla completelywhite; tube 21.2–22.5 mm long, 1.8–2.7 mmwide, 1.25–1.50 times as long as lobes,18.3–18.8 times as long as calyx, pubescent inupper half of tube inside, glabrous outside;lobes 15–18 × 4.7–6.2 mm, 2.9–3.2 times aslong as wide, elliptic or obovate, apex obtuse to
114 HARVARD PAPERS IN BOTANY Vol. 9, No. 1
acute, not ciliate, pubescent at very base oflobes inside, glabrous outside. Stamensinserted 18–19 mm from corolla base, which isca. 0.8 of corolla tube length in the rehydratedflowers measured; anthers 2.0–2.1 × 0.5 mm,4.0–4.2 times as long as wide, apex 2.2–2.3mm from corolla throat. Filaments 0.3–0.7 mmlong. Disk 0.9–1.0 mm long, 0.7–0.8 times aslong as ovaries, glabrous, awl-shaped, apexacute. Ovaries 1.1–1.4 mm high, glabrous orvery sparsely short pubescent on top; style16.3–16.4 mm long; style head 0.8–1.2 mmlong. Fruit oblique ellipsoid with an angle onone side, ca. 19 × 9 mm.
Distribution: China (Hainan)Habitat: in forest or on river banks from 240
to 430 m altitude.Additional collections studied: China:
Hainan: sin. loc. Katsumada s.n. (UC), Liang64643 (K, NY), Wang 33949 (A, NY);Yaichow, How 70734 (E); Po-ting, How 72659(A, BISH, BM, G, MO, P, PE), Fung 20018 (A,K, NY, UC, US); Bak Sa, Lau 27556 (A); DungKa, Chun & Tso 43853 (A, NY).
Typification: Tsiang gives the location of thetype specimens as in the herbarium ofSunyatsen University. These specimens arenow to be found at IBSC. Therefore the lecto-type has to be chosen from amongst the IBSC
duplicates of How 70532 and Chun & Tso43853, superceding Sévenet et al.’s designationof the CANT duplicate of How 70532 (Sévenetet. al., 1994).
E. D. Merrill has written the name Kopsiastenophylla Merr. n. sp. on Fung 20018 fromHainan, but there is no evidence that this namewas ever published.
10. Kopsia harmandiana [Pierre in L.Planch.,Prod. Apoc. 325. 1894, nom. nud.] ex Pitard,Fl. Gén. Indo-Chine 3: 1132. 1933; Ly, FeddesRepert. 97: 440. 1986; Sévenet et al., J.Ethnopharmacol. 41 (1994) 155. TYPE: VIET-NAM. Between the Mekong and Hué,Harmand in Pierre 5246 (Lectotype: P, desig-nated by Sévenet et al. [1994]; Isolectotypes:BR, P). Syntypes: P.A. Eberhardt 2740 (P), P.A.Eberhardt 2698 (P). Map 9.
Tree to 14 m tall. Branchlets glabrous,sparsely lenticellate. Leaves: petiole 2–8 mmlong, glabrous; blade 2.9–11.2 × 0.6–3.2 cm,2.1–6.7 times as long as wide, papery, subcori-aceous to coriaceous, elliptic or narrowly so,apex caudate to short acuminate with a blunttip, base cuneate, glabrous above and beneath,midrib raised and with a central groove above,flattened or shallowly sunken above, secondaryveins 16–31 pairs with 1.5–5 mm spacing,
MAP 9. Distribution of Kopsia harmandiana Pierre ex Pitard (•) and Kopsia hainanensis (♦).
60–75˚ from midrib, slightly prominent but notclearly distinguishable from tertiary venationabove and beneath, straight, tertiary venationprominent above and beneath, parallel to sec-ondary veins and reticulate, intramarginal veinright at margin. Inflorescence dichasial,2.5–5.5 cm long, branches 0.7 mm wide,glabrous; peduncles 2.3–3.5 cm long, 0.7 mmwide, glabrous; pedicels 1.5–2.0 mm long,glabrous, bracts present on pedicel. Sepals1.3–2.1 × 0.5–0.9 mm, 1.4–4.2 times as long aswide, lanceolate, apex acuminate, ciliate,glabrous outside, puberulent at tips inside.Corolla completely white; tube 15–23 mmlong, 1.7–2.1 mm wide, ca. 1.4 times as long aslobes, 10.0–11.5 times as long as calyx, pubes-cent in upper part of tube above, around, andslightly below the stamens and in throat,glabrous outside; lobes 13–17 × 4.5–6.2 mm,2.7–2.9 times as long as wide, obovate, apexacute, not ciliate, glabrous or pubescent at verybase of lobes inside, glabrous outside. Stamensinserted 8.0–9.7 mm from corolla base, whichis ca. 0.5 of corolla tube length in the rehy-drated flowers measured; anthers ca. 2.7 × 0.5mm, ca. 5.4 times as long as wide, apex 4.2–7.7mm from corolla throat; filaments 0.4 mmlong, pubescent. Disk 0.8–1.0 mm long, ca. 1times as long as ovaries, glabrous, awl-shapedor lanceolate, apex acute to acuminate. Ovaries0.8–1.0 mm high, densely pubescent on top;style 8.5 mm long; style head 1.1 mm long.Fruit falcate with a small spur, ca. 12 × 8 mm,spur 2 mm long, sparsely puberulent.
Distribution: Vietnam.Habitat: not recorded.Additional collections studied: Vietnam:
The location of the type is given vaguely asbetween the Mekong and Hué. This locality hasnot been included on the map, but wherever itactually is would be rather to the west of theother location marked on the map.
11. Kopsia lapidilecta [Sleesen, Fl. Mal. Misc.Rec. 1: 13. 1959, no Latin description] exSleesen, Blumea 10: 137. 1960; Markgr.,Blumea 20: 423. 1973; Sévenet et al., J.Ethnopharmacol. 41: 156. 1994; I.M.Turner,Gard. Bull. Sing. 47: 127. 1995. TYPE:INDONESIA. Natuna Islands, Gunung Ranai,C.G.G.J. van Steenis 1384 (Holotype: L;Isotype: BO). Map 7.
Habit unknown. Branchlets glabrous, denselylenticellate, terete. Leaves: petiole 3–8 mmlong, glabrous; blade 6.0–15.2 × 3.0–6.1 cm,2.0–2.6 times as long as wide, coriaceous,elliptic, apex short acuminate with a blunt tip,base obtuse, glabrous above and beneath,midrib flat to shallowly sunken or raised andwith a central groove above, secondary veins21–33 pairs with 2–6 mm spacing, 75–85˚ frommidrib, prominent above and beneath, notclearly distinguishable from tertiary venationabove, partly distinguishable or not from ter-tiary venation beneath, straight, tertiary vena-tion prominent above and beneath, parallel tosecondary veins or reticulate, intramarginalvein straight or only weakly looped, inset frommargin. Inflorescence with few elongatebranches, 3.5–7.0 cm long with axes 1.0–4.7cm long and branches 2.3 mm wide, glabrous;peduncle 0.2–0.5 cm long, 2 mm wide,glabrous; pedicels 1.3–2.0 mm long, glabrous;subtending bracts persistent; bracts present orabsent on pedicels. Sepals 1.7–2.0 × 1.4–1.5mm, 1.2–1.3 times as long as wide, ovate, apexrounded, ciliate, otherwise glabrous outsideand inside. Corolla bright red; tube 15–17 mmlong, 1.6–1.7 mm wide, 1.3–1.5 times as longas lobes, 7.5–10.0 times as long as calyx,densely pubescent beneath stamens and becom-ing less so further down, throat glabrous,glabrous outside; lobes 11.0–11.5 × 2.2–2.3mm, 4.8–5.2 times as long as wide, narrowlyoblong, apex obtuse, not ciliate, glabrous, out-side and inside. Stamens inserted 16.7–17.0mm from corolla base, which is ca. 0.8 ofcorolla tube length in the rehydrated flowersmeasured; anthers 2.2–2.4 × 0.5–0.6 mm,4.0–4.4 times as long as wide, apex ca. 0.9 mmfrom corolla throat; filaments ca. 1 mm long,glabrous. Disk ca. 1.4 mm long, 1.4–1.6 timesas long as ovaries, glabrous, awl-shaped, apexacute. Ovaries 0.9–1.0 mm high, glabrous;style ca. 16 mm long; style head ca. 0.7 mmlong. Fruit not seen (see comment below).
Distribution: Indonesia (Natuna Islands).Habitat: recorded from 200 m altitude.Additional collection studied: INDONE-
SIA. Kalimantan Barat, Bunguran, GunungRanai, 1385 (BO, L, SING).
Typification: Timmerman-Van der Sleesen isnot entirely clear about the typification of thisspecies, which under Art. 37.1 of theInternational Code of Botanical Nomenclaturewould only be validly published by reference to
116 HARVARD PAPERS IN BOTANY Vol. 9, No. 1
a single type specimen. She has written “Typus:Van Steenis 1384 (L); do 1385; Henderson SF20395. Anambas & Natuna Islands.” This couldbe interpreted as indicating that Van Steenis1384 in Leiden is the holotype or that all threeare syntypes. Only the former interpretationwould make it a validly published species.However, Markgraf (1973) attempted to typifythe species with Van Steenis 1385, which, ifallowed, would actually invalidate the publica-tion of the species. A type label has been placedon Van Steenis 1385 in Leiden, but the typeddetermination label by Timmerman-Van derSleesen also has a handwritten note saying“paratype,” possibly in her hand. Sévenet et al.(1994) gave the “holotype” as Van Steenis 1384(L), which, for the sake of convenience, I shallalso accept as a holotype, this being the inter-pretation of the original publication that wouldeffect no change. Henderson SF 20395, the onlyone of the paratypes from the Anambas Islandsrather than the Natuna Islands, has now beenmoved to the new species Kopsia grandifolia.
A fruit is illustrated in an accompanying illus-tration to the original description. However, noneof the original material now extant has any signof a fruit on it. The illustration shows a falcatefruit with only a small projection on one side.
12. Kopsia larutensis King & Gamble, J. As.Soc. Beng. 74(2): 432. 1907; Ridl., Fl. Mal.Pen. 2: 337. 1923; Sleesen, Fl. Mal. Misc. Rec.1: 13. 1959; Whitmore, Tree Fl. Mal. 2: 19.1972; Markgr., Blumea 20: 423. 1973; Sévenetet al., J. Ethnopharmacol. 41: 157. 1994;I.M.Turner, Gard. Bull. Sing. 47: 127. 1995.TYPE: Malaysia, Perak, Larut, L. Wray 2736(Lectotype: CAL, designated by Markgraf[1973: 423], first step, and here, second step[see comment below]; Isolectotypes: CAL,SING). Syntypes: King’s Collector 4269 (BM,K, L, SING), King’s Collector 6165 (BO, L,SING), King’s Collector 462 (K, P), King’sCollector 2082 (CAL), L. Wray 3956 (notfound), B. Scortechini 59b (BM, K), B.Scortechini 1704 (CAL), B. Scortechini 57b(BM). Map 10.
Tree to 1.8 m tall. Branchlets glabrous, notlenticellate, weakly angled. Leaves: petiole5–12 mm long, glabrous; blade 7.5–21.0 ×2.8–8.1 cm, 2.0–4.1 times as long as wide, sub-coriaceous to coriaceous, elliptic or oblong,apex caudate or more rarely long acuminate, inboth cases with a blunt tip, base acute tocuneate, glabrous above and beneath, midribshallowly sunken to raised above, or raised andwith a central groove, secondary veins 11–27
MAP 10. Distribution of Kopsia larutensis King & Gamble.
pairs with 4–11 mm spacing, 55–80˚ frommidrib, prominent to shallowly sunken above,prominent beneath, clearly distinguishablefrom tertiary venation above and beneath,curved ascending from midrib or only nearmargins, tertiary venation prominent above,prominent or not beneath, subperpendicular tomidrib and oblique to secondary veins and thensomewhat reticulate, intramarginal veinstraight or only weakly looped, inset from mar-gin. Inflorescence often many times branchednear the base with short thick branches thatappear very crowded, 1.7–6.0 cm long withaxes 0.4–5.2 cm long and branches 0.7–2.0 mmwide, glabrous or puberulent in upper parts;peduncle 0.3–5.0 cm long, 2.2 mm wide,glabrous; pedicels 1.0–1.5 mm long, glabrousor sparsely puberulent; subtending bracts per-sistent; bracts present on pedicel. Sepals1.3–2.1 × 1.2–1.8 mm, 1.1–1.5 times as long aswide, ovate, apex rounded, ciliate, glabrous orsparsely puberulent outside, glabrous inside.Corolla completely white; tube 7.0–10.5 mmlong, 1.1–1.3 mm wide, 1.3–1.4 times as longas lobes, 3.3–6.5 times as long as calyx, pubes-cent in lower half of tube but not at base, throatglabrous, glabrous outside; lobes 5–10 ×1.0–1.5 mm, 3.7–6.7 times as long as wide, lin-ear, apex acute, not ciliate, glabrous outside andinside. Stamens inserted ca. 6.5 mm fromcorolla base, which is ca. 0.8 of corolla tubelength in the rehydrated flowers measured;anthers ca. 1.4 × 0.4 mm, ca. 3.5 times as longas wide, apex ca. 0.3 mm from corolla throat,filaments 0.5 mm long, glabrous. Disk 0.7–1.1mm long, 0.7–1.0 times as long as ovaries,glabrous, awl-shaped, apex acute. Ovaries0.9–1.1 mm high, glabrous or very sparselypubescent on top; style ca. 5 mm long; stylehead ca. 0.7 mm long. Fruit with a blunthooked spur, 10–12 × 4.0–7.5 × 8–9 mm, spurca. 4 mm long, sparsely puberulent.
Distribution: Peninsular Malaysia (Perak).Habitat: understory shrub in lowland forest
to 200 m altitude.Selection of the 20 collections studied:
Typification: King and Gamble listed severalsyntypes for this species. A first stage lectotyp-ification in accordance with the St. Louis Code,article 9.14 (Greuter et al., 2000) was performedby Markgraf (1973). Sévenet et al. (1994)referred to a “holotype” in CAL but did not dis-tinguish between the two specimens of this col-lection to be found there. Therefore, this wasnot an effective second step lectotypification.
This is a very characteristic species withinflorescences that are branched near the basefollowed by crowded short cincinnate brancheswith small flowers that have narrow corollalobes.
The description given above will beexpanded when further and better collectionsare made of this species. Although there aremany collections, only Ng FRI 6024 had suffi-ciently mature and plentiful flowers to dissect,so it should be borne in mind that the androe-cium and gynoecium characters are based onmeasurements from this specimen only.
13. Kopsia macrophylla Hook.f., Fl. Brit. Ind.3: 639. 1882; King & Gamble, J. As. Soc.Beng. 74(2): 434. 1907; Ridl., Fl. Mal. Pen. 2:338. 1923; Sleesen, Fl. Mal. Misc. Rec. 1: 7.1959; Whitmore, Tree Fl. Mal. 2: 19. 1972;Markgr., Blumea 20: 423. 1973; Sévenet et al.,J. Ethnopharmacol. 41: 159. 1994; I.M.Turner,Gard. Bull. Sing. 47: 127. 1995. TYPE:Singapore, T. Lobb s.n. (Holotype: K; Isotype:BM). Map 11.Synonym: Kopsia ridleyana King & Gamble, J.
Tree to 6.1 m tall. Branchlets glabrous todensely puberulent, sparsely lenticellate or not,terete to weakly angled. Leaves: petiole 3–8mm long, glabrous to densely pubescent; blade7.5–24.5 × 1.6–9.2 cm, 2.2–4.6 times as long aswide, papery or subcoriaceous, elliptic to obo-vate, apex caudate or short to long acuminatewith a blunt tip, base acute or cuneate, glabrousto puberulent on midrib and major veinsbeneath, midrib shallowly sunken, flat or raisedand with a central groove above, secondaryveins 8–23 pairs with 3.3–15.0 mm spacing,50–65˚ from midrib, prominent or flat above,prominent beneath, clearly distinguishablefrom tertiary venation above and beneath,straight or curved ascending from midrib or
118 HARVARD PAPERS IN BOTANY Vol. 9, No. 1
only near margin, tertiary venation prominentabove and beneath, more or less parallel to sec-ondary veins or subperpendicular to midrib andoblique to secondary veins and then also some-what reticulate, intramarginal vein straight tostrongly looped, inset from margin, glabrous orpuberulent on midrib above. Inflorescencedichasial, mostly also with cincinnate branches,3.2–11.0 cm long with axes 0.8–7.3 cm longand branches 1.2–2.5 mm wide, glabrous todensely puberulent; peduncle 0.3–2.7 cm long,1.4–2.8 mm wide, glabrous to puberulent;pedicels 1.0–1.5 mm long, glabrous to denselypuberulent; subtending bracts persistent; bractspresent on pedicel. Sepals 1.7–2.5 × 1.6–1.7mm, 1.0–1.6 times as long as wide, ovate oroblong, apex rounded, margin ciliate, glabrousto densely puberulent, glabrous inside. Corollacompletely white or white with a yellow “eye”;tube 22–32 mm long, 1.4–1.9 mm wide,1.5–1.8 times as long as lobes, 8.8–16.0 timesas long as calyx, pubescent around stamens andslightly beneath inside or pubescent around andbeneath stamens and in throat, glabrous in-between, throat pubescent, sometimes soshortly so as to appear papillose, glabrous out-side; lobes 13–21 × 5.3–12.5 mm, 1.2–2.5times as long as wide, elliptic or broadly ellip-tic, apex rounded, ciliate or not, glabrous out-
side and inside. Stamens 8.5–12.0 mm fromcorolla base, which is 0.3–0.5 of corolla tubelength in the rehydrated flowers measured;anthers 2.3–2.4 × 0.4–0.7 mm, 3.4–5.1 times aslong as wide, apex 9.7–12.5 mm from corollathroat; filaments 0.6 mm long. Disk 0.6–1.3mm long, 0.6–1.1 times as long as ovaries,glabrous, awl-shaped, apex acute to obtuse.Ovaries 0.8–1.2 mm high, sparsely to denselypubescent on top; style 8.5–9.6 mm long; stylehead ca. 1 mm long. Fruit with a blunt hookedspur from about the middle of the fruit, 15–18× 8.0–8.5 mm, spur ca. 4 mm long, glabrous orsparsely puberulent.
Distribution: Peninsular Malaysia.Habitat: Recorded from 100 to 152 m altitude.Selection of the 14 collections studied:
MALAYSIA: Johor: Gunong Angsi, Ridley11903 (K); Between Perhentian Tinggi andGunong Angsi, Ernst 1074 (L). Melaka, Alvins2178 (SING). Negri Sembilan: Seremban,Bukit Tangga, Napier s.n. (SING). Pahang:Dong, near Raub, Burkill & Haniff 16904(SING); Bukit Serdam, Henderson 25044 (K,NY, SING); Rompin, Mahamud 17183 (KEP,SING), Watson CF3194 (K, KEP, SING);Lawang Ulu Lung, Mat Soh 15426 (K, KEP).Perak: sin. loc. King’s Collector 4963 (K).SINGAPORE, T. Lobb s.n. (BM, K).
MAP 11. Distribution of Kopsia macrophylla Hook.f.
Typification: Sévenet et al. (1994) typifiedKopsia macrophylla with King 4963 fromPerak, despite the fact that this collection wasmade in 1883, a year after the species was pub-lished. Hooker had noted that he had seen onlyone specimen collected by Lobb in Singapore.Lobb s.n. in Kew is, therefore, the holotype.
Kopsia ridleyana was first included in syn-onymy of Kopsia macrophylla by Timmerman-Van der Sleesen (1959), and was followed byMarkgraf (1973) and Sévenet et al. (1994).Timmerman-Van der Sleesen noted that the leafcharacters, which, in part, were used by Kingand Gamble (1907) to distinguish the twospecies, break down with the larger number ofspecimens available since Kopsia ridleyanawas described. However, there are differencesbetween typical K. macrophylla and typical K.ridleyana. Kopsia macrophylla has a denselypubescent inflorescence and is sparsely todensely pubescent on the twigs and on theundersurface of the leaf blades. Kopsia rid-leyana is glabrous and also tends to havesmaller flowers. However, there are specimensthat have a pubescent inflorescence butglabrous twigs and leaves, and specimens withsmall flowers but pubescent inflorescences,leaves, and twigs. These bridge the gapbetween the extremes and so prevent a demar-cation of taxa. The variation in this species israther akin to that found in K. pauciflora var.pauciflora. Of the form with densely pubescentinflorescences there appear to be two subforms:one with a distinctly pubescent corolla throatand another where the hairs in the throat appearto be so short as to appear more or less papil-lose. The first form is throughout the range ofthe species, the latter from Pahang inPeninsular Malaysia. There are otherwise nodifferences in other characters. No new mater-ial of either form has been collected since 1960.
14. Kopsia pauciflora Hook.f., Fl. Brit. Ind. 3:639. 1882; King & Gamble, J. As. Soc. Beng.74(2): 431. 1907; Ridl., Fl. Mal. Pen. 2: 337.1923; Sleesen, Fl. Mal. Misc. Rec. 1: 14. 1959;Whitmore, Tree Fl. Mal. 2: 19. 1972; Markgr.,Blumea 20: 422. 1973; Anderson, ChecklistTrees Sarawak 149. 1980; P.S.Ashton, ManualNon-dipt. Trees Sarawak 38. 1988; Sévenet etal., J. Ethnopharmacol. 41: 160. 1994;I.M.Turner, Gard. Bull. Sing. 47: 127. 1995.TYPE: MALAYSIA. Malacca: Mount Ophir,
A.C. Maingay KD 1056 (Holotype: K; Isotypes:K, L [scrap]; Photograph: A). Maps 12–13.Synonyms: Kopsia parvifolia Merr., Phil. J.
Sci. 29: 412. 1926; Masam., Enum. Phan.Born. 621. 1942; Sleesen, Fl. Mal. Misc.Rec. 1: 8. 1959; Sévenet et al., J.Ethnopharmacol. 41: 160. 1994; Beamanet al., Plants Mount Kinabalu 4: 108.2001. TYPE: MALAYSIA. Sabah:Banguey Island [=Pulau Banggi], A.Castro & F. Melegrito 1456 (Holotype:UC; Isotypes: A, BM, BO, K).
MAP 12. Distribution of Kopsia pauciflora Hook.f. var. pauciflora.
MAP 13. Distribution of Kopsia pauciflora Hook.f. var. mitrephora (Sleesen) D.J.Middleton.
Kopsia pauciflora Hook.f. var. pauciflora. Map 12.
Tree to 10 m tall; to 15 cm dbh. Bark gray,olive-brown or white, smooth; inner bark palebrown, straw colored or white. Branchletsglabrous to sparsely puberulent, lenticellate ornot, angled, weakly winged or markedlywinged. Leaves: petiole 0–10 mm long,glabrous; blade 5.0–25.5 × 1.1–9.5 cm, 1.8–6.1times as long as wide, subcoriaceous to coria-ceous, elliptic, apex caudate or short to longacuminate with a blunt tip, base rounded tocuneate, glabrous or puberulent on midribabove, glabrous or puberulent beneath, midribdeeply to shallowly sunken above, or raisedabove, sometimes then with a central groove,secondary veins 7–24 pairs with 2–19 mmspacing, 55–80˚ from midrib, prominent or flatabove, prominent beneath, clearly distinguish-able or not from tertiary venation above andbeneath, straight or curved ascending frommidrib or only near margin, tertiary venationprominent or flat above and beneath, some-times rather obscure, parallel or subperpendic-ular to midrib and oblique to secondary veins,then also somewhat reticulate, intramarginalvein obscure, straight or only weakly looped.Inflorescence dichasial or with cincinnatebranches, 3.5–14.0 cm long with axes 0.5–10.0cm long and branches 0.3–3.1 mm wide,glabrous to densely puberulent; peduncle0.0–1.8 cm long, 1.2–3.2 mm wide, glabrous todensely puberulent; pedicels 0.5–2.5 mm long,glabrous to densely puberulent; subtendingbracts persistent; bracts present on pedicel.Sepals 1.5–2.9 × 1.0–3.2 mm, 0.8–2.0 times aslong as wide, ovate or oblong, apex rounded,ciliate, glabrous to densely puberulent outside,glabrous or puberulent on upper half inside.Corolla completely white, white with yellow“eye,” white and green or, rarely, pinkish-white; tube 25–44 mm long, 1.6–2.8 mm wide,1.2–2.4 times as long as lobes, 10.9–19.5 timesas long as calyx, pubescent around stamens andslightly beneath inside, throat pubescent,glabrous or sparsely puberulent at top of tubeoutside; lobes 11.5–33.0 × 4.6–19.0 mm,1.7–4.1 times as long as wide, elliptic or obo-vate, apex rounded, ciliate or not, glabrous out-side and inside. Stamens 22–33 mm fromcorolla base which is 0.7–0.9 of corolla tubelength in the rehydrated flowers measured;anthers 1.8–2.8 × 0.4–0.8 mm, 2.6–5.6 times aslong as wide, apex 2.4–5.0 mm from corollathroat; filaments 0.5–0.8 mm long, glabrous or
pubescent. Disk 0.9–1.9 mm long, 0.8–1.9times as long as ovaries, glabrous, awl-shapedor deltoid, apex acute, acuminate or irregularlytoothed. Ovaries 0.9–1.5 mm high, sparsely todensely pubescent, or mostly glabrous with afew hairs; style 20.0–31.5 mm long; style head1.0–1.3 mm long. Fruit falcate with a blunthooked spur from about the middle of the fruit,13.5–17.0 × 4.5–6.0 × 6–9 mm, hook 3–5 mmlong, sparsely puberulent.
Distribution: Southern Thailand, PeninsularMalaysia, Sumatra, Borneo, Java (but see notebelow).
Habitat: in a wide variety of forest types, ona wide variety of soil types, from sea level to1500 m altitude.
Geographical selection of the 165 collec-tions studied: BRUNEI DARUSSALAM:River Batu Apoi–Sebatu watershed, AshtonBRUN310 (K, L). INDONESIA: Java: Cicurug,Jampango, Backer 17199 (K, L, SING). NatunaIslands: West of Kampong Ranai, Steenis 1097(L). Sumatra: Riau, Bünnemeyer 5962 (L).MALAYSIA: Terengganu: Sungai Kerbat atJeram Petang, Whitmore FRI20351 (SING, K,L, A, KEP). Perak: Piah Forest Reserve, Jaamat89235 (SING). Sabah: Mt. Silam, 12 milesWSW of Lahad Datu, Beaman 10053 (GH, K,L, NY); Kota Merudu, Bukit Mandalong, AminGambating SAN109817 (K, L, SAN). Sarawak:Maputi, Brooke 10159 (L), 10164 (BM, G, L,US), 10185 (BM, L). THAILAND: Krabi: AoLuk, Charoenphol, Larsen & Warncke 3446 (L,P). Phangnga: Sanangmanora Forest Park,Middleton 570 (A, BKF). Narathiwat: BachoNational Park, Kiah 24285 (K).
Typification: Sévenet et al. (1994) desig-nated the BM duplicate of A. Castro & F.Melegrito 1456 the lectotype (but incorrectlyused the term holotype) of Kopsia parvifolia.However, Merrill quite clearly states in the pro-tologue that the still extant UC duplicate is theholotype. Similarly, Sévenet et al. (1994) des-ignated the NY duplicates of A.D.E. Elmer20130 and A.D.E. Elmer 20615 the “holotypes”(= lectotypes) of K. caudata and K. caudatavar. glabra, respectively. Merrill, however, inan introduction to the paper in which thesespecies were described, notes that he based hisdescriptions on his examination of the materialin PNH and UC, and therefore the lectotypesmust be chosen from amongst this material.Given that the PNH material has beendestroyed, the UC material must be lectotypi-fied for both taxa.
122 HARVARD PAPERS IN BOTANY Vol. 9, No. 1
Only one collection, Backer 17199, is knownfrom Java. Although it is not indicated on thelabel, this collection may have been taken froma cultivated plant and so the species may not benative to Java.
The plant is described variously as a shrub orsmall tree on almost all specimens except forMadani et al. SAN142633, on which it isdescribed as a 70 m tall tree with a 50-m clearbole and a girth of 60 cm. This is such an enor-mous tree (albeit with an unlikely girth) and soout of keeping with all the other material of thisspecies, and of the genus as a whole, that I sus-pect a mistake has been made and so the dimen-sions are not included in the description above.
This is an extremely variable species both invegetative and floral morphology, although thisvariation is mostly manifest in the Borneanspecimens rather than the continental ones. Thefruit, however, is fairly uniform. The patterns ofvariation are reticulate. At the extremes, thespecimens with thickly coriaceous leaves andanther apices only a few millimetres below thethroat of the corolla tube appear very differentfrom the specimens with thinner leaves and sta-mens inserted in the lower half of the corollatube. However, around the Telupid area inSabah specimens with particularly and charac-teristically thick leaves and a fairly uniformvegetative morphology show almost the entirerange of stamen position within the tube.Beaman 10241 and Gardner 79, both fromTawai in Sabah, have leaves with a very shortpetiole but would otherwise appear to fallwithin the variation of Kopsia pauciflora.There are also specimens scattered elsewherewith stamens inserted high in the corolla tubeand thin leaves or stamens inserted low in thecorolla tube and thick leaves. There are, how-ever, relatively few specimens that are reallyintermediate in stamen position, leading tointriguing and unanswered questions about thepollination biology of this species.
Larsen et al. 43442 from Ao Luk in Krabi inThailand has a label describing the flower coloras pinkish white. It is the only specimen not tohave the flower color described as either whiteor white with a yellow “eye” but is otherwiseunremarkable for Kopsia pauciflora. In the areawhere it was collected, the newly described andundoubtedly closely related K. rosea, whichhas either white or pink flowers, is also found,and the possibility arises that there has beensome hybridization between the two taxa.
10: 136. 1960; Sleesen, Fl. Mal. Misc.Rec. 1: 7. 1959; Whitmore, Tree Fl. Mal.2: 19. 1972; Markgr., Blumea 20: 424.1973; Sévenet et al., J. Ethnopharmacol.41: 159. 1994. TYPE: MALAYSIA.Sabah: Lahad Datu District, Pathbetween Sungei Sabahan and SungeiDok, G.H.S. Wood SAN16118 (Holotype:L; Isotype: BRI).
Tree to 3 m tall. Branchlets glabrous,sparsely lenticellate or not, weakly angled tomarkedly winged. Leaves: petiole 2–7 mmlong, glabrous; blade 5–24 × 1.5–7.5 cm,2.4–5.1 times as long as wide, subcoriaceous orcoriaceous, elliptic or oblong, apex long acumi-nate with a blunt tip, base acute to cuneate,glabrous above and beneath, midrib shallowlysunken to raised above, secondary veins 13–22pairs with 3–14 mm spacing, 65–75˚ frommidrib, prominent above and beneath, clearlydistinguishable from tertiary venation aboveand beneath, straight to curved ascending frommidrib, tertiary venation prominent above andbeneath, parallel to secondary veins or subper-pendicular to midrib and oblique to secondaryveins, usually also somewhat reticulate, intra-marginal vein straight or looped, inset frommargin. Inflorescence dichasial, 4.3–6.5 cmlong with axes 0.9–3.4 cm long and branches0.8–1.9 mm wide, glabrous, peduncle 0.2–0.8cm long, 0.8–2.0 mm wide, glabrous, pedicels0.7–2.5 mm long, glabrous or sparsely puberu-lent; subtending bracts persistent; bracts pre-sent on pedicel. Sepals 1.3–2.5 × 1.1–1.6 mmwide, 1.25–1.80 times as long as wide, ovate oroblong, apex rounded, ciliate or not, glabrousoutside and inside. Corolla completely white orwhite with a yellow “eye,” tube 15.0–28.5 mmlong, 1.1–1.5 mm wide, 0.8–2.0 times as longas lobes, 10.7–13.6 times as long as calyx,pubescent around stamens and slightly beneathinside, throat pubescent, glabrous outside;lobes 9–28 × 3.5–11.5 mm, 2.0–3.1 times aslong as wide, elliptic, apex rounded to obtuse,ciliate or not, glabrous outside and inside.Stamens inserted 3.5–15.5 mm from corollabase, which is 0.1–0.6 of corolla tube length inthe rehydrated flowers measured; anthers1.7–2.3 × 0.4–0.6 mm, 3.4–5.5 times as long aswide, apex 6.5–19.0 mm from corolla throat,
filaments 0.5–0.8 mm long, pubescent. Disk0.7–1.4 mm long, 0.7–1.6 times as long asovaries, glabrous, awl-shaped, apex acute toacuminate. Ovaries 0.5–1.2 mm high, sparselyto densely pubescent; style 2.5–12.5 mm long;style head 0.8–1.3 mm long. Fruit falcate witha small blunt hooked spur on one side, 12–18 ×2.5–4.5 × 6.0–7.5 mm, hook 3–5 mm long,sparsely puberulent.
Distribution: Malaysia (Sabah).Habitat: in primary forest.Selection of the 29 collections studied:
1995. TYPE: MALAYSIA. Terengganu,Dungun, C. Wiart & L.E. Teo KL 4432(Holotype: P; Isotypes: KEP, P).
Tree to 3 m tall. Bark gray. Branchletsglabrous, sparsely to densely lenticellate or not.Leaves: petiole 2–12 mm long, glabrous; blade5.2–22.0 × 1.4–6.8 cm, 3.1–4.4 times as long aswide, papery, elliptic or oblong, apex caudateto long acuminate with a blunt tip, basecuneate, glabrous above and beneath, midribshallowly sunken, to raised and and then some-times with a central groove above, secondaryveins 12–20 pairs with 2.5–18.0 mm spacing,50–70˚ from midrib, prominent above andbeneath, clearly distinguishable from tertiaryvenation above and beneath, ascending nearmargin or curved ascending from midrib, ter-tiary venation prominent above and beneath,parallel to secondary veins or irregularly sub-perpendicular to midrib and oblique to sec-ondary veins, intramarginal vein straight oronly weakly looped, inset from or right at mar-gin. Inflorescence dichasial or cincinnate,3.5–14.0 cm long with axes 1.7–12.0 cm longand branches 0.9–1.5 mm wide, glabrous;peduncle 0.8–5.4 cm long, 1.0–1.8 mm wide,glabrous; pedicels 1.2–3.5 mm long, glabrousor sparsely puberulent, subtending bracts
124 HARVARD PAPERS IN BOTANY Vol. 9, No. 1
MAP 14. Distribution of Kopsia profunda Markgr.
deciduous, bracts present on pedicel. Sepals1.0–2.1 × 1.0–1.4 mm, 0.8–1.5 times as long aswide, ovate, apex obtuse to acute, ciliate,glabrous outside, glabrous or puberulent at tipsinside. Corolla completely white or white witha yellow “eye”; tube 20–26 mm long, 1.1–6.1mm wide, 1.3–1.8 times as long as lobes,10.2–21.0 times as long as calyx, pubescent inupper part of tube above or around and slightlybelow the stamens, pubescent in throat,glabrous outside; lobes 11.5–20.0 × 3.8–8.0mm, 1.8–3.7 times as long as wide, elliptic,apex obtuse to acute, ciliate or not, glabrousoutside and inside. Stamens inserted 6.0–9.1mm from corolla base, which is 0.3–0.4 ofcorolla tube length in the rehydrated flowersmeasured; anthers 1.7–2.5 × 0.4–0.6 mm wide,3.8–5.0 times as long as wide, apex 1.1–10.7mm from corolla throat; filaments 0.6–1.0 mmlong, pubescent. Disk 0.7–1.1 mm long,0.8–1.1 times as long as ovaries, glabrous, awl-shaped, apex acute. Ovaries 0.8–1.1 mm high,densely long pubescent on top or denselypubescent all over; style 4.9–7.3 mm long; stylehead 0.9–1.5 mm long. Fruit falcate with asmall or blunt hooked spur, ca. 1.8 × 6 mm,hook 4 mm long, sparsely puberulent.
Distribution: Peninsular Malaysia.Habitat: in lowland evergreen forest from
230 to 330 m altitude.Additional collections studied: Terengganu:
Bukit Kajang, Corner 25936 (K, SING); UluBrang, Moysey & Kiah SFN.33892 (K, L,SING); Dungun, Hutan Simpan Bukit Bauk,Anthony 604 (KEP), Teo & Pachiapu 1080 (K,L, SING), Sevenet & Deverre 242 (P), Deverre135 (P), Teo, Pachiapu & Boichard 1186 (P);Dungun, David 217 (P), Hutan Simpan BukitKesing, Teo & Pachiapu 1136 (K, L, SING);49–50 km from Dungun to Kuala Berang, Teo& Din KL 5035 (P).
The only differences between this speciesand Kopsia griffithii are in the stamens, whichare borne lower in the corolla tube, and the lackof pubescence on the disk. They share the intra-marginal vein, the yellow-eyed corolla, theacute sepals, and the lax inflorescence.
Sévenet et al. (1994) distinguished Kopsiaterengganensis from K. profunda on the posi-tion of the stamens in the corolla tube. Theysuggested that K. profunda had stamens situ-ated at the base of the tube and K. terengga-nensis in the middle. In fact, the typespecimens of K. profunda and of K. terengga-nensis have stamens situated at exactly a third
of the length of the corolla tube from the base.They are indistinguishable in other characters.
Sévenet et al. (1994) suggested that Kopsiaterengganensis (= K. profunda) differed fromK. macrophylla in being erect as opposed toprostrate. However, the only specimen of eitherspecies I have seen where the habit is describedas prostrate was Sévenet & Deverre 242, whichwas incorrectly labelled as K. macrophylla butis in fact K. profunda. The leaves are also justas large in this specimen as in K. macrophylla.However, in K. profunda the inflorescence isdelicate, lax, and few-flowered and the sepalsacute, characters which do suggest that it is dis-tinct from K. macrophylla.
Kopsia larutensis auct. non King &Gamble: Anderson, Checklist TreesSarawak 149. 1980; P.S.Ashton, ManualNon–dipt. Trees Sarawak 38. 1988.
Kopsia arborea auct. non Blume:Markgr., Blumea 20: 419. 1973, pro parte.
Small tree or shrub, to 4.6 m tall. Branchletsglabrous, not or sparsely lenticellate, terete.Leaves: petiole 4–9 mm long, glabrous; blade13.4–32.0 × 3.8–9.6 cm, 2.6–4.3 times as longas wide, papery to subcoriaceous, elliptic oroblong, apex caudate, base obtuse to cuneate,glabrous above and beneath, midrib shallowlysunken or raised and with a central grooveabove, secondary veins 9–25 pairs with 6–25mm spacing, 40–60˚ from midrib, prominent orflat above, prominent beneath, clearly distin-guishable from tertiary venation above andbeneath, straight or slightly ascending nearmargin, tertiary venation prominent above,prominent or flat beneath, reticulate or sub-per-pendicular to midrib and oblique to secondaryveins, intramarginal vein looped and inset frommargin. Inflorescences dichasial and then withcincinnate branches, 4–15 cm long with axes1.4–20.0 cm long and branches 1.4–2.1 mmwide, glabrous to densely puberulent; peduncle0.2–7.6 cm long, 1.7–3.7 mm wide; pedicelsca. 4 mm long; subtending bracts persistent.Sepals 1.5–1.7 × 1.1–1.4 mm, 1.2–1.4 times as
long as wide, ovate, apex rounded, ciliate, oth-erwise glabrous outside and inside. Corollawhite; tube 21.5–26.5 mm long, ca. 2.3 mmwide, pubescent around stamens and slightlybeneath inside, throat pubescent, glabrous out-side, 1.8–2.0 times as long as lobes, 12.6–16.7times as long as calyx; lobes 11–15 × 2.7–4.7mm wide, 3.0–4.5 times as long as wide, ellip-tic or oblong, apex obtuse or acute, not ciliate,glabrous outside and inside. Stamens 17–21mm from corolla base which is ca. 0.8 ofcorolla tube length in the rehydrated flowersmeasured; anthers 1.7–2.0 × 0.6–0.8 mm,2.5–2.8 times as long as wide, apex 1.1–2.0mm from corolla throat; filaments ca. 0.7 mmlong. Disk ca. 0.9 mm high, ca. 1 times as longas ovaries, glabrous, awl-shaped, apex acute.Ovaries ca. 0.9 mm high, glabrous to denselypubescent; style ca. 20 mm long; style head ca.0.8 mm long. Fruits falcate with small bluntprojection near the base, 15–16 × 4 × 6.5–7.0mm; projection 3–4 mm long.
Distribution: Malaysia (Sarawak).Habitat: in primary or disturbed forest to
300 m altitude.Additional collections studied: MALAYSIA:
Sarawak: Kapit, Upper Rajang River, Clemens& Clemens 21211 (K); Pelagus Rapids, AshtonS.17797 (L); Rajang, Haviland 3042 (BM, K,
SAR, SING); Bukit Raya, Smith S.27738(SAR).
This recently described species is knownfrom only a few specimens from Sarawak and,as can be seen from the citations, has long beenconfused with a number of other species. It ismost immediately recognizable by its largeleaves and the angle of the secondary veinswith the midrib, and inflorescences that canbecome very long.
17. Kopsia rosea D.J.Middleton, sp. nov.TYPE: THAILAND. Krabi, Ao Luk, ThanbokKhorani National Park, D.D. Soejarto, T.Santisuk, K. Taylor & N. Nantasan 5945(Holotype: A; Isotypes: F, L, MO, NY). Map 16.
Frutex ad 3 m altus, ramulis glabris. Foliaelliptica, 4.7–27.5 × 2.3–11.0 cm, nervis 8–20paribus. Corollae tubus 24–45 mm longus,lobis 15–31 × 9.5–16.5 mm.Usage synonyms: Kopsia alba auct. non Ridl.
ex Henderson: Kerr, Fl. Siam. Enum. 2:437. 1939, pro parte.
Kopsia fruticosa auct. non (Roxb.)A.DC.: Middleton, Fl. Thailand 7: 61.1999, pro parte.
Kopsia macrophylla auct. non Hook.f.:Middleton, Fl. Thailand 7: 63. 1999.
126 HARVARD PAPERS IN BOTANY Vol. 9, No. 1
MAP 15. Distribution of Kopsia rajangensis D.J.Middleton (■), Kopsia sleeseniana Markgr. (•), and Kopsiatenuis Leenh. & Steenis (♦).
Tree to 10 m tall. Branchlets glabrous, lenti-cellate or not, terete or weakly angled. Leaves:petiole 3.0–12.5 mm long, glabrous; blade4.7–27.5 × 2.3–11.0 cm, 1.9–3.2 times as long aswide, coriaceous, elliptic, apex short acuminatewith a blunt tip, base acute or cuneate, glabrousabove and beneath, midrib shallowly sunkenabove, secondary veins 8–20 pairs with 6.5–18.0mm spacing, 60–70˚ from midrib, prominent orflat above, prominent beneath, clearly distin-guishable from tertiary venation above andbeneath, curved ascending from midrib or onlynear margin, tertiary venation prominent or flatabove and beneath, obscure or parallel or sub-perpendicular to midrib and oblique to sec-ondary veins, then also somewhat reticulate,intramarginal vein straight or only weaklylooped, inset from margin. Inflorescencedichasial, often with robust cincinnate branches,5.0–12.5 cm long with axes 1.0–9.5 cm long andbranches 1.4–2.9 mm wide, glabrous to denselypuberulent; peduncle 0.4–4.5 cm long, 1.3–3.0mm wide, glabrous or puberulent; pedicels0.0–6.5 mm long, glabrous to densely puberu-lent; subtending bracts persistent; bracts presenton pedicel. Sepals 1.5–3.5 × 1.4–2.2 mm,1.1–1.6 times as long as wide, oblong, apexrounded, ciliate, glabrous to densely puberulentoutside, glabrous inside. Corolla white, palepink with darker pink “eye,” or white with a pinktinge; tube 24–45 mm long, 2.0–2.7 mm wide,1.1–1.5 times as long as lobes, 8.3–16.0 times aslong as calyx, pubescent in upper part of tubeabove, around and slightly below the stamens,throat pubescent, glabrous or sparsely puberu-lent at top of tube outside; lobes 15–31 ×9.5–16.5 mm, 1.8–1.9 times as long as wide,obovate, apex rounded, not ciliate, glabrous out-side and inside. Stamens 22.4–35.0 mm fromcorolla base, which is ca. 0.8 of corolla tubelength in the rehydrated flowers measured;anthers 2.2–2.6 × 0.6–0.8 mm, 2.9–4.0 times aslong as wide, apex 2.2–4.0 mm from corollathroat; filaments 0.7–1.1 mm long, glabrous.Disk 0.8–2.0 mm long, 0.7–1.4 times as long asovaries, glabrous, lanceolate, apex acuminate.Ovaries 1.2–2.5 mm high, glabrous or with justone or two hairs; style ca. 22.5 mm long; stylehead ca. 1.2 mm long. Fruit falcate with a blunthooked spur from around the middle of the fruit,15–18 × 4.5–5.5 × 8–9 mm, spur 2.5–4.5 mmlong, sparsely puberulent.
This new species has similarities to Kopsiapauciflora, differing from it in the glabrousovaries and in mostly having a pink or pink-tinged corolla. The robust inflorescence struc-ture is reminiscent of K. macrophylla, fromwhich it differs in the stamens being insertedhigher up the corolla tube. It differs from thepink-flowered K. fruticosa in the glabrousovaries and in the glabrous twigs. Middleton(1999), working on the Thai species only,included the pink-flowered specimens of thisspecies in a more variable K. fruticosa and thewhite-flowered specimens in a more variableK. macrophylla. With a broader knowledge ofthe ranges of variation in characters in thegenus as a whole, it is now clear that K. fruti-cosa is native only in Burma and is morpholog-ically less variable than previously thought, andthat K. macrophylla is only in PeninsularMalaysia and contains only specimens with sta-mens inserted low down in the corolla tube.Kopsia fruticosa is, however, widely cultivatedin Thailand and could easily be mistaken for K.rosea, except on closer examination.
glabrous; blade 6.1–19.5 × 2.7–11.4 cm,1.4–2.8 times as long as wide, subcoriaceous tocoriaceous, elliptic, apex short acuminate withblunt or sharp tip, base acute or cuneate,glabrous above and beneath, midrib sunkenabove, secondary veins 7–15 pairs with 4–16 mmspacing, 45–80˚ from midrib, prominent above,prominent or flat beneath, clearly distinguish-able from tertiary venation above and beneath,curved ascending from midrib, tertiary vena-tion prominent above, prominent or flatbeneath, irregularly subperpendicular to midriband oblique to secondary veins, intramarginalvein straight or only weakly looped, inset fromor right at margin. Inflorescence dichasial,mostly many-flowered, 3.5–12.5 cm long withaxes (0.5–)1.2–11.5 cm long and branches1.2–1.8 mm wide, glabrous or puberulent inupper parts; peduncle (0.2–)1.0–7.5 cm long,1.4–1.8 mm wide, glabrous or puberulent;pedicels 1.0–5.3 mm long, glabrous to denselypuberulent, subtending bracts persistent, bractspresent or absent on pedicel. Sepals 2–4 ×1.4–3.4 mm, 1.0–1.8 times as long as wide,ovate or oblong, apex rounded or obtuse, cili-ate, glabrous to densely puberulent outside,glabrous inside. Corolla white with red “eye”;tube 32–36 mm long, 1.8–2.9 mm wide,
1.4–2.1 times as long as lobes, 8.75–16.50times as long as calyx, pubescent inside above,around and slightly below the stamens, throatpubescent, glabrous or rarely sparsely puberu-lent at top of tube outside; lobes 15–24 ×6.5–12.0 mm, 2.0–3.1 times as long as wide,elliptic or oblong, apex rounded to obtuse, cili-ate, glabrous or, rarely, sparsely pubescent inupper quarter outside, glabrous or puberulent atbase inside. Stamens inserted 23.5–34.0 mmfrom corolla base, which is 0.8–0.9 of corollatube length in the rehydrated flowers measured;anthers 1.7–2.1 × 0.7–0.8 mm, 2.4–3.0 times aslong as wide, apex 1.2–2.2 mm from corollathroat; filaments 0.2–1.0 mm long, pubescent.Disk 1.1–1.8 mm long, 0.6–1.1 times as long asovaries, glabrous, oblong or awl-shaped, apexacute to acuminate. Ovaries 1.6–1.9 mm high,sparsely to densely pubescent on top; style27–30 mm long; style head 1.1–1.2 mm long.Fruit oblique ellipsoid with either a simpleangle or a spur on one side, 26–33 × 6–7 ×11.5–14.0 mm, angle/spur 2.5–3.0 mm long,glabrous.
Distribution: Peninsular Malaysia,Singapore.
Habitat: in lowland evergreen forest, swampforest or on river banks to 600 m.
128 HARVARD PAPERS IN BOTANY Vol. 9, No. 1
MAP 16. Distribution of Kopsia rosea D.J.Middleton (■) and Kopsia singapurensis Ridl. (•).
FIGURE 7. Kopsia singapurensis Ridl. A, habit; B, flower dissection; C, ovaries and disk; D, fruit. From ShahMS.1226 (A, B, C) and David 263 (D). Scale bars = 1 cm (A, B, D); 1 mm (C).
Geographical selection of the 47 collec-tions studied: MALAYSIA. Johor: nearMersing, David 085 (P), 263 (P, A); SungaiKaya, Mawai-Jemulang Road, Corner 28731(A, BM, K, KEP, SING), Kiah 31959 (A, K,KEP, SING), 32039 (SING, A, BRI, KEP, BM).SINGAPORE: Seletar Reservoir, NoorSRMN17 (A, CANB, L, SING).
Typification: Ridley indicated a number ofsyntypes of this species in his original descrip-tion and indicated that his own collection wasfrom Singapore, Chan Chu Kang, but withoutgiving a collection number. Later, Sévenet et al.(1994) lectotypified the species with “Ridley59, BM!, 1892.” However, the only specimenlabelled as Ridley 59 I could find in BM wascollected in 1890 and this specimen wasreferred to Kopsia singapurensis by Ridleyhimself, unlike a number of the other speci-mens from this locality. Other specimens col-lected by Ridley from Chan Chu Kang in otherherbaria were collected in 1889, 1891, 1892,and 1894 and so may be syntypes although,again, most of them have not been labelled asK. singapurensis by Ridley. A specimen inKew, Ridley s.n. (collected in 1894), labelled asK. singapurensis by Ridley himself, is here des-ignated as the lectotype, superceding Sévenet etal.’s designation, which could not be locatedand was possibly misassigned.
Although King and Gamble (1907) veryclearly make the new combination Kopsia fru-ticosa var. albiflora (Teijsm. & Binn.) King &Gamble based on Calpicarpum albiflorumTeijsm. & Binn., it is clear from their com-ments and specimens cited that they are actu-ally refering to the plant later described as K.singapurensis. Calpicarpum albiflorum, thetype of which is from Seram, is now treated asa synonym of K. flavida.
This species is close to Kopsia flavida, bothhaving a white corolla with a red “eye,” and thefruit has a broad deltoid projection rather thenthe smaller, often hooked projection of many ofthe other species. However, K. singapurensishas a larger and laxer inflorescence, moreprominent tertiary venation, and a much lessprominent projection on the fruit.
19. Kopsia sleeseniana Markgr., Blumea 20:421. 1973; Sévenet et al., J. Ethnopharmacol.41: 161. 1994. TYPE: MALAYSIA. Sarawak,Bintulu, G.D. Haviland 3046 (Holotype:SING; Isotypes: K, L, SAR). Map 15.
Small tree reported at 2 m tall. Branchletsglabrous, not lenticellate, terete or weaklyangled. Leaves: petiole 7–12 mm long glabrous;blade 8.5–21.5 × 3.6–7.8 cm, 2.2–5.0 times aslong as wide, papery to subcoriaceous, ellipticor oblong, apex caudate or long acuminate witha blunt tip, base rounded to cuneate, glabrousabove and beneath, midrib shallowly sunken orraised and with a central groove above, sec-ondary veins 23–43 pairs with 2–9 mm spacing,70˚ from midrib, prominent above and beneath,weakly distinguishable from tertiary venationabove and beneath, straight or ascending nearmargin, with intercalcated weaker parallel ter-tiary veins, tertiary venation prominent aboveand beneath, intramarginal vein straight or onlyweakly looped, inset from or right at margin.Inflorescence dichasial, lax, robust, higher orderbranching is not opposite but alternate, 6.0–10.2cm long with axes 2.2–7.5 cm long andbranches 1.6–2.2 mm wide, glabrous; peduncle0.6–4.5 cm long, 1.4–2.2 mm wide, glabrous;pedicels 2.8–5.2 mm long, glabrous; subtendingbracts persistent; bracts absent on pedicel.Sepals 1.3–1.6 × 1.0–1.6 mm, 1.0–1.3 times aslong as wide, ovate, apex rounded or obtuse, cil-iate, glabrous outside and inside. Corolla com-pletely white; tube 26.5–32.0 mm long, 1.9–2.6mm wide, 1.6–2.1 times as long as lobes,18.7–20.4 times as long as calyx, pubescentaround stamens and slightly beneath inside,throat pubescent, glabrous outside; lobes13.2–20.0 × 3.3–6.0 mm, 3.3–4.0 times as longas wide, narrowly elliptic or oblong, apexobtuse, not ciliate, glabrous outside and inside.Stamens inserted ca. 22 mm from corolla base,which is ca. 0.8 of corolla tube length in therehydrated flowers measured; anthers ca. 2.4 ×0.8 mm, ca. 3 times as long as wide, apex 1.1mm from corolla throat; filaments ca. 0.9 mmlong, pubescent. Disk ca. 0.8 mm long, ca. 0.7times as long as ovaries, glabrous, awl-shaped,apex acuminate. Ovaries ca. 1.2 mm high,densely pubescent on top; style ca. 19.4 mmlong; style head ca. 0.9 mm long. Fruit flattenedoblique ellipsoid with a blunt hooked spur, ca.16–17 × 7 × 4 mm, densely puberulent.
Distribution: Malaysia (Sarawak)Habitat: lowland mixed dipterocarp forest
and beach forest to 80 m altitude.
130 HARVARD PAPERS IN BOTANY Vol. 9, No. 1
Additional collections studied: Sarawak:Bintulu, Similajau National Park, JawaS.65629 (SAR), S.65651 (SAR); Niah NationalPark, Sungai Sekaloh, Ching S.40133 (K, L,SAN, SAR).
The most distinctive feature of this species isthe lax and robust inflorescence. The holotypeis, unfortunately, in fairly poor condition.
20. Kopsia sumatrana D.J. Middleton, sp.nov. TYPE: INDONESIA. Sumatra: RiauProvince, Bukit Karampal, 5 km W ofTalanglakat on Rengat–Jambi Road, J.S.Burley, Tukirin et al. 1886 (Holotype: A;Isotype: L). Map 17; Fig. 8.
Frutex 2 m altus, ramulis puberulis. Foliaelliptica, oblonga vel ovata, 11.5–26.0 ×2.7–5.5 cm, nervis 16–19 paribus. Corollaetubus ca. 19.5 mm, lobis 14 × 1.7 mm.
Tree reported to 2 m tall, 2 cm dbh.Branchlets densely puberulent, becomingglabrescent with age, not lenticellate, weaklyangled. Leaves: petiole 4.0–7.5 mm long,densely pubescent; blade 11.5–26.0 × 2.7–5.5cm, 3.5–5.4 times as long as wide, papery,elliptic, oblong or ovate, apex acuminate withblunt tip, base obtuse to cuneate, puberulent onmidrib only or sparsely puberulent on midrib
and major veins above, puberulent on midriband major veins beneath, midrib shallowlysunken above, secondary veins 16–19 pairswith 3–13 mm spacing, ca. 60˚ from midrib,prominent above and beneath, not clearly dis-tinguishable from tertiary venation, ascendingnear margin, tertiary venation prominent aboveand beneath, irregularly parallel to secondaryveins, intramarginal vein straight or onlyweakly looped, inset from margin.Inflorescence cincinnate, 5.0–7.2 cm long withaxes 3.4–5.5 cm long and branches 1.4–1.7 mmwide, densely puberulent; peduncle 0–5 cmlong, 2.6 mm wide, puberulent; pedicels 1–2mm long, densely puberulent, subtendingbracts persistent, bracts present on pedicel.Sepals ca. 1.4 × 1.4 mm, ca. 1 times as long aswide, ovate, apex rounded, ciliate, denselypuberulent outside, glabrous inside. Corollayellowish; tube ca. 19.5 mm long in dried flow-ers, ca.1 mm wide, ca. 1.4 times as long aslobes, ca. 13.9 times as long as calyx, pubes-cent in upper part of tube above, around andslightly below the stamens inside, throatglabrous, sparsely puberulent at top of tube out-side; lobes ca. 14 × 1.7 mm, ca. 8.2 times aslong as wide, linear, obtuse, not ciliate,glabrous outside and inside. Stamens inserted
MAP 17. Distribution of Kopsia sumatrana D.J.Middleton (■) and Kopsia teoi L.Allorge (•).
132 HARVARD PAPERS IN BOTANY Vol. 9, No. 1
FIGURE 8. Kopsia sumatrana D.J.Middleton. A, habit; B, flower dissection; C, ovaries, disk, and base of style;D, close up of underside of leaf. From Turkirin et al. 1820 (A, B, C, D). Scale bars = 1 cm (A, B); 1 mm (C, D).
ca. 19.5 mm from corolla base in rehydratedflowers, which is ca. 0.8 of corolla tube lengthin the rehydrated flowers measured; anthers ca.2.5 × 0.5 mm, ca. 5 times as long as wide, apexca. 1.9 mm from corolla throat; filaments ca.0.7 mm long, pubescent. Disk 0.9 mm long, ca.0.7 times as long as ovaries, glabrous, awl-shaped or narrowly deltoid, apex acute toobtuse. Ovaries ca. 1.3 mm high, glabrous orsparsely pubescent on top; style ca. 18 mmlong; style head ca. 0.8 mm long.
Distribution: Sumatra.Habitat: recorded from ridge forest at 100 m
altitude.Additional collection studied: INDONE-
SIA: Sumatra Riau: Bukit Karampal, 5 km westof Talanglakat on Rengat–Jambi road, Burley etal. 1820 (A).
This new species is very distinctive with itscombination of pubescent leaves and extremelynarrow corolla lobes.
Tree or shrub to 6 m tall, to 15 cm dbh. Barkbrown, slightly flaky. Branchlets glabrous,sparsely lenticellate or not, terete. Leaves: peti-ole 1–2 mm long, glabrous; blade 3.2–13.7 ×1.2–4.5 cm, 2.2–4.6 times as long as wide,papery, elliptic, oblong or ovate, apex caudate,base acute to cuneate, glabrous above andbeneath, midrib shallowly sunken above, sec-ondary veins 16–30 pairs with 1–3 mm spac-ing, 70–80˚ from midrib, prominent above, flatbeneath, not clearly distinguishable from ter-tiary venation above or beneath, straight, ter-tiary venation prominent or flat above, flatbeneath, obscure or clearly visible, mostly par-allel to secondary veins, intramarginal veinright at margin. Inflorescence delicate, lax,dichasial or flowers solitary, 3.7–17.0 cm longwith axes 1.1–15.0 cm long and branches0.5–5.0 mm wide, sparsely puberulent; pedun-cle 0.2–2.2 cm long, 0.4–5.0 mm wide, puberu-lent, pedicels 4.5–5.0 mm long, glabrous orsparsely puberulent, subtending bracts persis-tent, bracts normally 2 on pedicel. Sepals 1 ×0.9–1.0 mm, 1.0–1.1 times as long as wide,
ovate, rounded, ciliate, glabrous. Corolla com-pletely white or yellowish; tube 16–17 mmlong, 1.8 mm wide, 1.1–1.4 times as long aslobes, 16–17 times as long as calyx, pubescentaround stamens and slightly beneath inside,throat glabrous or with a few sparse hairs,glabrous outside; lobes 11.5–15.0 × 2.4–3.0mm, 3.80–6.25 times as long as wide, narrowlyelliptic or oblong, apex obtuse, not ciliate,glabrous outside and inside. Stamens insertedca. 14 mm from corolla base, which is ca. 0.8of corolla tube length in the rehydrated flowersmeasured; anthers ca. 2 × 0.4 mm, ca. 5 timesas long as wide, apex ca. 2 mm from corollathroat; filaments ca. 0.4 mm long, pubescent.Disk ca. 0.6 mm long, ca. 0.7 times as long asovaries, glabrous, awl-shaped, apex obtuse.Ovaries ca. 0.9 mm high, glabrous; style ca. 13mm long; style head ca. 0.9 mm long. Fruit flat-tened oblique ellipsoid with a blunt hookedspur, 12.5–15.0 × 5.6–7.3 × 3.0–3.4 mm, spur3–4 mm long, arising in lower part of fruit,sparsely puberulent (see comments on thisdescription below).
Distribution: Malaysia (Sarawak)Habitat: recorded from mixed dipterocarp
forest on sandy clay soils from 20 to 735 m alti-tude.
Collections studied: Sarawak: sin. loc.Beccari 1861 (K, P); Mt. Poi (=Gunong Pueh),Clemens & Clemens 20129 (A, BO, K, L, NY,SAR, Z), 20178 (SAR), Mjoberg s.n. (UC),Paie 13625 (K, L, SAR), Purseglove P.4819(A, K, L, SAR, SING), James et al. S.34459(A, K, L, MO, SAN, SAR); Mount Matang,Clemens & Clemens 20898 (K, MO, NY,SAR), Ridley s.n. (K); Semunsam WildlifeSanctuary, Ching S.43441 (K, KEP, L, SAN,SAR).
The inflorescence structure is quite variable,with between 1 and 6 flowers in an inflores-cence. A note on the type specimen, which hasonly 2 flowers of quite different size, suggeststhat the larger flower is abnormal. However, thelarger flower corresponds far more to the flow-ers of all other specimens of this species, so thedescription given here does not correspond tothe original description with measurements thatare taken from the smaller flower, which,although badly damaged and difficult to inter-pret, may simply be immature.
FIGURE 9. Kopsia tenuis Leenh. & Steenis. A, habit; B, flower dissection; C, ovaries and disk. From James etal. S.34459 (A, B, C). Scale bars = 1 cm (A, B); 1 mm (C).
127. 1995. TYPE: Malaysia, Johor, Keluang,L.E. Teo & F. Remy KL 3976 (Holotype: P;Isotypes: KEP, KLU, n.v.). Map 17.
Tree to 4 m tall. Branchlets glabrous,sparsely lenticellate or not, terete. Leaves ses-sile; blade 6–22 × 2.1–9.2 cm, 2.1–3.2 times aslong as wide, subcoriaceous or coriaceous,elliptic, apex short acuminate with a blunt tip,base rounded to acute, glabrous above andbeneath, midrib shallowly sunken to raisedabove, sometimes also with a central groove,secondary veins 11–15 pairs with 5–18 mmspacing, 50–70˚ from midrib, prominent aboveand beneath, clearly distinguishable from ter-tiary venation above and beneath, curvedascending from midrib, tertiary venationprominent above and beneath, subperpendicu-lar to midrib and oblique to secondary veins, orobliquely scalariform, also somewhat reticu-late, intramarginal vein straight to stronglylooped, inset from margin. Inflorescencedichasial, 5.6–6.3 cm long with axes 0.3–1.7cm long and branches 0.7–1.3 mm wide,glabrous; peduncle 0.0–1.2 cm long, 1.4–1.7mm wide, glabrous; pedicels 1.2–2.5 mm long,glabrous; subtending bracts persistent; bractsseveral on each pedicel. Sepals 1.6–2.9 ×1.3–2.0 mm, 1.1–1.5 times as long as wide,ovate, apex rounded to acute, ciliate, glabrousoutside, puberulent at tips inside. Corolla whitewith a pink tinge or pink with a red “eye”; tube34–41 mm long, 2.7–2.8 mm wide, 1.6–2.3times as long as lobes, 14.1–25.6 times as longas calyx, pubescent in upper part of tube above,around, and slightly below the stamens, throatpubescent, glabrous outside; lobes 18–25 ×8–14 mm, 1.3–2.2 times as long as wide, ellip-tic or broadly elliptic, apex rounded or obtuse,not ciliate, glabrous outside and inside.Stamens inserted 35.0–37.5 mm from corollabase, which is ca. 0.9 of corolla tube length inthe rehydrated flowers measured; anthers1.7–1.9 × 0.5 mm, 3.4–3.8 times as long aswide, 1.8–1.9 mm from corolla throat; fila-ments 0.9–1.4 mm long, glabrous. Disk ca. 1.2mm long, ca. 1.5 times as long as ovaries,glabrous, awl-shaped, apex acute. Ovaries ca.0.8 mm high, glabrous or sparsely pubescent ontop; style 34.0–37.5 mm long; style head0.9–1.1 mm long. Fruit unknown.
Distribution: Peninsular Malaysia.Habitat: in lowland Dipterocarp forest to
200 m altitude.Additonal collections studied: MALAYSIA:
Johor: Ulu Madik, Holttum 10634 (BM);
Endau–Rompin State Park, Sungai Semawak,near Nature Education Centre, Saw FRI44947(KEP); Kwala Kahang, Lake & Kelsall s.n.(SING); Mersing, Teo & Remy KL4018 (KEP);Taman Negara Endau, Sungai Selai, Teo & DinKL 5011 (P).
The most distinctive features of this speciesare the sessile leaves, constant in this speciesbut never so in other species, and the severalbracts on each pedicel.
23. Kopsia tonkinensis Pitard, Fl. Gén. Indo-Chine 3: 1135. 1933; Ly, Feddes Repert. 97:440. 1986; Sévenet et al., J. Ethnopharmacol.41: 162. 1994. TYPE: VIETNAM. Phu-ThoProvince, between Thanh-ba and Chan-mongForest Reserve, F. Fleury 32111 (Holotype: P;Isotype: P). Map 18.
Tree to 12 m tall. Branchlets sparsely todensely puberulent, especially when young,sparsely lenticellate or not, weakly angled.Leaves: petiole 7–15 mm long, glabrous; blade7.3–21.1 × 2.2–8.3 cm, 2.1–3.9 times as long aswide, subcoriaceous to coriaceous, elliptic orovate, apex acuminate with blunt tip, baseobtuse to cuneate, glabrous above and beneath,midrib sunken above, secondary veins 12–19pairs with 4–22 mm spacing, (45–)60–70˚ frommidrib, prominent beneath, prominent or flatabove, clearly distinguishable from tertiaryvenation, ascending near margin, tertiary vena-tion prominent above and beneath, reticulate orsubperpendicular to midrib and oblique to sec-ondary veins, intramarginal vein straight,weakly or quite strongly looped, inset frommargin, sometimes not evident at all.Inflorescence dichasial or cincinnate, 4.1–11.2cm long with axes 1.0–7.5 cm long andbranches 1.1–1.7 mm wide, glabrous orsparsely puberulent; peduncle 0.1–3.7 cm long,1.4–1.7 mm wide, glabrous or puberulent;pedicels 0.5–2.3 mm long, glabrous, subtend-ing bracts deciduous or persistent, bracts pre-sent or absent on pedicel. Sepals 1.3–2.0 ×1.0–1.8 mm, 1.0–1.8 times as long as wide,ovate or oblong, apex rounded or emarginate,ciliate, glabrous outside and inside. Corollacompletely white, pinkish-white, or brown-red;tube 21–31 mm long, 1.6–3.3 mm wide,1.5–2.3 times as long as lobes, 10.5–18.5 timesas long as calyx, pubescent around and beneathstamens and in throat, glabrous outside; lobes9–17 × 4.5–7.5 mm, 1.8–2.5 times as long aswide, elliptic or obovate, apex rounded, ciliateor not, glabrous outside and inside. Stamens
inserted 17–21 mm from corolla base, which is0.7–0.8 of corolla tube length in the rehydratedflowers measured; anthers 2.0–2.5 × 0.50–0.65mm, 3.8–5.0 times as long as wide, apex2.0–4.5 mm from corolla throat; filaments0.5–1.0 mm long, pubescent. Disk 0.6–1.0 mmlong, 0.7–0.9 times as long as ovaries,glabrous, awl-shaped, apex acute or acuminate.Ovaries 0.9–1.3 mm high, glabrous; style15–21 mm long; style head 1.00–1.75 mmlong. Fruit (only immature seen) falcate withsmall blunt hooked spur, sparsely puberulent.
Distribution: Vietnam.Habitat: recorded from evergreen forest at
300 m altitude.Additional collections studied: VIETNAM:
District, Chau Kim, Ban Muong, Soejarto et al.10203 (GH).
Typification: Although there are three speci-mens of Fleury 32111 in the Paris herbarium,only one of them has detailed notes of a dissec-tion attached and Kopsia tonkinensis written onthe label. The other two specimens are unana-lyzed and without an indication that theyformed part of original material. Therefore, thespecimen with the dissection notes is taken tobe the holotype rather than needing to be lecto-typified.
Close to Kopsia fruticosa, differing in the sta-mens being slightly lower in the corolla tubeand in the glabrous ovaries. As for most speciesthere is a woeful lack of information providedon the labels. Three specimens provide infor-mation on the color of the corolla: one says thecorolla is white, another pinkish-white, and theother that it is brown-red. This third specimenlacks corollas altogether on the specimen, butthe inflorescence structure fits the pattern forthe species.
136 HARVARD PAPERS IN BOTANY Vol. 9, No. 1
MAP 18. Distribution of Kopsia tonkinensis Pitard.
Poilane 6650 from near Nhatrang in Vietnamhas only extremely immature flowers butwould appear to be an undescribed Kopsiaspecies. Most characteristic for the plant is the
very long petioles, up to 2 cm long, longer thanfor any other species in the genus. It also has aglabrous, lax inflorescence.
Kopsia majumdarii M.G.Gangop. & Chakrab.,J. Econ. Tax. Bot. 16: 59. 1992. The holotype of this species, King’s Collector
(= Kunstler) 7937 (CAL), is a specimen of thesame collection as was used to describe Kayeacaudata King in the Clusiaceae, now treated asa synonym of Kayea kunstleri King, althoughthe specimen, plus two isotype specimens inCAL, do not bear any indication that Kingintended these particular specimens as part ofthe original material of Kayea caudata. PeterStevens has seen these specimens and confirmsthey are Kayea kunstleri in the Clusiaceae.
EXCLUDED SPECIES
Kopsia elastica Pierre in L.Planch., Prod.Apoc. 325. 1894, nom. nud.
Kopsia parallela Regel & Koern., Ind. Sem.Hort. Petrop. 44. 1857.
There is insufficient information to identifythis species and no specimen could be traced bythe St. Petersburg herbarium staff.
INSUFFICIENTLY KNOWN
ALLORGE, L. 1993. Kopsia teoi L.Allorge(Apocynaceae), a new Malayan species. Acta Bot.Gallica 140: 97–99.
ALLORGE, L., and L. E. Teo. 1986. A new Kopsiafrom Malaysia (Apocynaceae). Phytologia 59:93–94.
BLUME, C. L. 1823. Catalogus. Batavia.———. 1826. Bijdragen tot de Flora van
Nederlandsch Indië. Lands Drukkerij, Batavia.———. 1849. Rumphia. Vol. 4. C. G. Sulpke,
Leiden.BRUMMITT, R. K., AND C. E. POWELL. 1992. Authors
of plant names. Royal Botanic Gardens, Kew.DE CANDOLLE, A. L. L. P. 1844. Prodromus. Vol. 8.
Treuttel & Würtz, Paris.DON, G. 1837. Apocyneae. A General History of the
Dichlamydeous Plants 4: 69–105. Rivington et al.,London.
ENDRESS, M. E., AND P. V. BRUYNS. 2000. A revisedclassification of the Apocynaceae s.l. BotanicalReview 66(1): 1–56
FORMAN, L. L. 1997. Notes concerning the typifica-tion of names of William Roxburgh’s species ofphanerogams. Kew Bulletin 52: 513–534
GREUTER, W., J. MCNEILL, F. R. BARRIE, H. M.BURDET, V. DEMOULIN, T. S. FILGUEIRAS, D. H.NICOLSON, P. C. SILVA, J. E. SKOG, P. TREHANE, N.J. TURLAND, AND D. L. HAWKSWORTH, EDS. 2000.International Code of Botanical Nomenclature(Saint Louis Code) adopted by the SixteenthInternational Botanical Congress. St. Louis,Missouri, July–August 1999. Koeltz, Königstein.
HOLMGREN, P. K., N. H. HOLMGREN, AND L. C.BARNETT. 1990. Index Herbariorum. IAPT, NewYork.
HOOKER, J. D. 1882. Flora of British India. Vol. 2.Reeve and Co., London.
KER GAWLER, J. B. 1819. Cerbera fruticosa. Bot.Reg. 5: t.391.
KING, G., AND J. S. GAMBLE. 1907. Materials for aflora of the Malayan Peninsula. Apocynaceae. J.Roy. Asiat. Soc. Bengal 74(2): 387–505.
LI, P. T., A. J. M. LEEUWENBERG, AND D. J.MIDDLETON. 1995. Apocynaceae. In Flora ofChina 16: 143–188. Science Press, Beijing.
MARKGRAF, F. 1973 [‘1972’]. Florae MalesianaePraecursores LIII. Apocynaceae II. 6. Urnularia,7. Willughbeia, 8. Kopsia. Blumea 20: 407–425.
LITERATURE CITED
———. AND K. HUBER. 1975. Die Entwicklung derNasenfrucht von Kopsia flavida Bl. BotanischeJahrbücher 96: 256–269.
MERRILL, E. D. 1926. The flora of Banguey Island.Philipp. J. Sci. 29: 341–427.
———. 1929. Plantae Elmerianae Borneenses.University of California Publications in Botany15: 1–316.
MERRILL, E. D., and L. M. PERRY. 1941. A summaryof Kentrochrosia Lauterbach and Schumann.Philipp. J. Sci. 76: 19–21.
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———. 2003. A new species and a new combinationin Bornean Kopsia (Apocynaceae). Gard. Bull.Singapore 55: 65–68.
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H. HUMBERT, EDS., Flore Géneral de l’Indochine 3: 1087–1262.
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SÉVENET, T., L. ALLORGE, B. DAVID, K. AWANG, A.HAMID, A. HADI, C. KAN–FAN, J.-C. QUIRION, F.REMY, H. SCHALLER, AND L. E. TEO. 1994. A pre-liminary chemotaxonomic review of Kopsia(Apocynaceae). J. Ethnopharmacology 41:147–183.
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The number after each collection refers to the number given to the species above except for 8a =Kopsia griffithii var. griffithii, 8b = Kopsia griffithii var. pubescens, 14a = Kopsia pauciflora var. pau-ciflora, 14b = Kopsia pauciflora var. mitrephora. Only specimens with a clearly identified collectorand collection number are given.
The number following each name is the number of the species under which a name can be found,except for the following: (g) see under genus heading, (x) excluded species, and (ik) insufficientlyknown. Recognized taxa are in roman type face, synonyms are italicized, and new taxa are in boldface.
A previously undescribed species of Kopsia from Vietnam has been identified but, unfortunately,too late to be added to this revision. It will be described shortly. It is most similar to K. grandifoliaand K. sumatrana in the narrow corolla lobes but differs from both in the glabrous branchlets andsepals and the lax inflorescence.
![E-ISSN: Triterpenes and sterols of family …It is one of the important latex-forming families [3]. Apocynaceae is divided into five subfamilies: Rauvolfioideae, Apocynoideae, Periplocoideae,](https://static.documents.pub/doc/80x56/5e6884c63d966b79b724168a/e-issn-triterpenes-and-sterols-of-family-it-is-one-of-the-important-latex-forming.jpg)
