revision of disepalum (annonaceae)

Brittonia, 41(4), 1989, pp. 356-378. �9 1989, by the New York Botanical Garden, Bronx, NY 10458-5126

R E V I S I O N O F D I S E P A L U M ( A N N O N A C E A E )

DAVID M. JOHNSON

Johnson, David M. (Department of Botany-Microbiology, Ohio Wesleyan Uni- versity, Delaware, OH 43015). Revision of Disepalum (Annonaceae). Brittonia 41: 356-378. 1989.--The relationship between the Asian generaDisepalum Hook. f. and Enicosanthellum Ban (Annonaceae) is examined in a revisionary study. Despite the disparate perianth morphologies of the two genera, other characters support inclusion of all species in a single genus with nine species. The monocarp stalks in all species are found to be derived from the receptacle rather than the ovary, and are thus not homologous with those found in most other Annonaceae genera. Cladistic analysis indicates that the species on the mainland of Asia and of montane habitats are more primitive than the species of the lowlands and of Sumatra and Borneo. Two new combinations, Disepalum petelotii and D. plagioneurum, are necessary as a result of merging the two genera, and one species, D. acieulare, is described as new.

Disepalum Hook. f. is a small genus of trees and shrubs indigenous to lowland and mon tane forests o f southeastern Asia. I ts m e m b e r s are peculiar a m o n g An- nonaceae in having flowers with two rather than three sepals. The flowers exhibit characteristics o f petal morpho logy remarkable for the family, including petals connate in a single whorl, petal n u m b e r that regularly varies f rom four to nine, and special modif icat ions o f the petal apices (Fig. 1 D--J). Previous keys to species of Disepalum (van Steenis, 1948; Fries, 1959) have emphas ized petal n u m b e r in distinguishing species, but it is now clear that these t rea tments do not account adequately for var ia t ion in this character.

Three species originally described in the large paleotropical genus Polyalthia Blume, P. pulchra King, P. petelotii Merrill, and P. plagioneura Diels, have been proposed for inclusion in Disepalum, despite their t r imerous perianths with free petals: Sinclair (1955)just i f ied his p lacement o f P. pulchra in Disepalum on the basis o f similarities in s tamens and carpels, and Walker (1971) found that P. plagioneura 1 shared with Disepalum species large tectate-columellate pollen in octads. Ban (1975) erected a new genus, Enicosanthellum Ban, to a c c o m m o d a t e the latter two species, and P. pulchra was later t ransferred to it by van Heusden (in Maas et al., 1988), who commented , "Th i s t r a n s f e r . , , leaves Disepalum again in its fo rmer c i rcumscr ipt ion as a very natural- looking genus."

This study is part o f a larger project to evaluate the tribe Bocageae Endl. (="Tr igynaea G r u p p e " of Fries, 1959; = " C y m b o p e t a l u m Tr ibe" of Walker , 1971) of the Annonaceae, to which Walker (1971) assigned Disepalum. In order to evaluate Walker ' s hypothesis , as well as to identify consistent differences for distinguishing Disepalum taxa, and to address the p rob lem of how widely the genus is to be circumscribed, I examined he rba r ium specimens of all m e m b e r s o f the genus, including mater ia l o f the species that have been assigned to Eni- cosanthellum.

Systematic Criteria

Phyllotaxy and Architecture. - -Phyl lo taxy on lateral shoots of mos t Annonaceae is distichous, and exceptions to this pat tern in the family are o f potent ial phy-

In fact, Walker suggested that both P. plagioneura and P. petelotii should be transferred to Dise- palum on the basis of his examination of the pollen of the vouchers, How 70331 and Pdtelot 6362, respectively. Both of these specimens, however, are P. plagioneura.

1989] JOHNSON: DISEPALUM 357

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FIo. 1. Representative flowers of the species of Disepalum, showing variation in perianth morphology. A. D. petelotii (P~telot 1780, NY). B. D. plagioneurum (P~telot 6362, NY). C. D. pulchrum (Anderson 2949, MO). D. D. platypetalum (Bangham & Bangham 1232, A); note sutures between petals, and intervening regions of receptacle without attached petals (black areas). E. D. platypetalum (de Wilde & de Wilde-Duy~es 18353, L); petals without sutures and entirely adnate to receptacle. F. D. coronatum (Sibat ak Luang S.17148, L), four-petaled flower. G. D. coronatum (Beaman 8064, NY), six-petaled flower. H. D. aciculare (Anderson 13118, SING). I. D. longipes (Bremer & Bremer 1826, KLU). J. D. anomalum (van Niel 4102, MO).

logenetic significance. Reports o f 2/5 phyllotaxy observed in Disepa lum a n o m - a lum Hook. f. (Fries, 19 5 9; Koek -Noorman et al., 19 8 8) are therefore of interest. In all material o f Disepa lum and of species assigned to Enicosan the l lum that I examined, however, the phyllotaxy of the lateral shoots was strictly distichous. I

358 BRITTONIA [VOL. 41

found, upon re-examination of the literature, that both reports of 2/5 phyllotaxy stemmed from a herbarium study of the construction of flowering shoots of D. anomalum by Wagner (1906). In his speculation concerning the architecture of the Disepalum tree, Wagner suggested that in D. anomalum the older growth might have some condition, for example 2/5 phyllotaxy, other than the distichy found on the flowering shoots he studied, observing that if the phyllotaxy were distichous throughout the individual, a peculiar flattened plant would result.

Hall6 et al. (1978) categorized most Annonaceae as conforming to one of two architectural models, the principal distinction between them being that in Roux's Model the trunk is monopodial and orthotropic, bearing plagiotropic lateral branches, while in Troll's Model the "trunk" is composed of the basal portions of successively superposed plagiotropic axes. Wagner thus anticipated Roux's Model, an architecture known to be exhibited by at least seven genera of Annon- aceae (Hall6 et al., 1978); Disepalum anomalum, if it has such an orthotropic trunk, would therefore not be unusual in the family. It does not, however, have orthotropy in the lateral shoots, and comparisons to genera of Annonaceae that do have orthotropic lateral shoots, such as Tetrameranthus R. E. Fries, are not valid.

Inflorescences and Flowers.- Flowers of all species in the study group are borne terminally on ebracteate pedicels. If the leaf opposing the pedicel is reduced in size and replacement shoot growth has not begun, the inflorescence appears bracteate and truly terminal. If replacement shoot growth has begun, the pedicel appears simply leaf-opposed (Murray, 1988). Inflorescences consist of a single flower in the species that have been assigned to Enicosanthellum and in Disepalum platypetalum Merr., and of one or two flowers in D. coronatum Becc., D. aciculare D. M. Johnson, D. longipes King, and D. anomalum.

The sepals of all species in the study group are foliaceous, valvate in aestivation, and are large for the family. They enclose the bud but eventually become reflexed.

The petals in the species that have been assigned to Enicosanthellum are relatively large (Fig. 1 A-C), completely free to the base, and are usually imbricate at the apices in the young stages. In Disepalum sens. strict., the petals are uniformly smaller (Fig. 1D-J), imbricate at the apices in young buds, and are not only connate into a short broad tube, but this tube is also adnate for most of its length to the broad convex receptacle. An intermediate condition between the free petals of the species assigned to Enicosanthellum and the adnate and connate petals of Disepalum sens. strict, is found, however, in some specimens ofD. platypetalum, where an individual flower may have both free and connate petals. In addition, D. platypetalum has petals that are shaped like those in the species that have been assigned to Enicosanthellum, and are intermediate in size between the two groups (Fig. 1D, E).

Stamens and Pollen.- All species have hundreds of clavate stamens with cushion-shaped connectives and short filaments crowded on the broad receptacle of each flower (Fig. 1 E, G, H, J). In D. coronatum, D. aciculare, and D. longipes the connectives become bright red at anthesis, making the pendent flowers very conspicuous from below. Walker (1971) described the pollen of Disepalum in detail. He found the genus to be uniformly characterized by pollen consisting of large tectate-columellate grains in octads, and suggested the transfer of Polyalthia plagioneura to Disepalum on this basis. I found the pollen of Enicosanthellum pulchrum and E. petelotii, species not examined by Walker, to have the same morphology. The polyads in the undehisced anthers are arranged in columns within each theca, which gives the appearance of partitions between the polyads. Despite a report by Ban (1975), however, I could find no septa present in the mature stamens.

1989] JOHNSON" DISEPALUM 359

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FIG. 2. Monocarps and carpophores in species of Disepalum, and in representative species of Neostenanthera, Enantia, and Polyalthia. A. D. petelotii (Henry 10886, NY). B. D. plagioneurum (Ting & Shih 1086, L). C. D. pulchrum (Stone 15371, KLU). D. D. platypetalum (de Wilde & de Wilde-DuyJ}es 18622, L). E. D. coronatum (Beaman 8064, NY). F. D. aciculare (Sinclair & bin Tassim 10193, US). G. D. longipes (Shah & Ahmad MS 2916, SING). H. D. anomalum (Corner BR UN 5350, SING). I. Neostenanthera harnata (Benth.) Exell (de Wilde 3349, K). J. Enantia lebrunii Robyns & Ghesq. (Lebrun 4231, NY). K. Polyalthia rumphii (BI.) Merr. (Sinclair & bin Tassim 10444, NY).

Carpe l s . - -Carpe l s vary in number f rom ca 200 per flower in several species to ca 18 in D. peteloti i . The carpels are club-shaped and pubescent, with the style and ovary subequal in length until anthesis, at which t ime the style apex expands greatly. The one or two (rarely three) ovules are attached laterally (if ovule solitary, this condit ion may be termed sub-basal by some authors), usually in the lower half of the carpel locule, not basally at tached as has been reported in the literature (e.g., Fries, 1959; Walker, 1971; Morawetz, 1986).

F r u i t s � 9 fruits of both D i s e p a l u m sens. strict, and o f the species that have been assigned to E n i c o s a n t h e l l u m are superficially similar to those found in many genera of Annonaceae, consisting o f an aggregate of stipitate, fleshy, few-seeded, indehiscent monocarps. The remarkable feature of the fruits in all species o f the study group is that the stalks bearing the monocarps develop not from the base of the ovary, but f rom the receptacular tissue immediately below it. This is manifested in an articulation between the monocarp and its stalk (Fig. 2), and in the texture of the stalk resembling that o f the torus rather than that o f the monocarp. The stalks are therefore termed carpophores rather than monocarp stipes in this paper. I have found similar carpophores to occur elsewhere in the family in the African genera N e o s t e n a n t h e r a Exell and E n a n t i a Oliver (Fig. 2).

S e e d s . - - C h r i s t m a n n (1986) described the seed coats o f D i s e p a l u m (Enicosan-


thellum) pulchrum and D. anomalum as consisting of inner and outer layers, with oil idioblasts occurring in the nucellus. This type of seed coat histology was also found by Christmann in the seeds of species in 10 other genera, however, so it is not decisive concerning the relationship between Enicosanthellum and Disepalum. The seed coat in all species of the study group was smooth, shiny, and not adherent to the carpel wall, with a conspicuous papery rim to the exostome. There is no vestige of an aril or aril-like structure.

Summary.--The species of Disepalum sens. strict, and Enicosanthellum share many similarities in both vegetative and reproductive characters, although the difference in perianth morphology between the two genera cannot be disputed. An intermediate floral morphology does, however, occur; the example of Dise- palum platypetalum previously mentioned, in which the petals are not always connate and are similar in shape to those found in Enicosanthellum, with the entire flower larger than that of most other species of Disepalum sens. strict., provides a link between the two extremes.

Sympetaly is a highly significant taxonomic character in many flowering plant groups, but in the Annonaceae it is a phenomenon that reappears in diverse evolutionary lines. Sympetalous species occur within otherwise cohesive poly- petalous genera such as Annona L., Fusaea (Baill.) Saff., and Uvaria L. (Fries, 1959), Popowia Endl. sens. strict. (Sinclair, 1955), and Polyalthia (pers. obs.).

Similarly, in many groups difference in number of perianth parts is important taxonomically. Although dimerous flowers occur among otherwise trimerous flowers of Annonaceae as teratologies, in several taxa within otherwise trimerous genera, dimery has become fixed: several species ofDesmos Lour. (sensu Sinclair, 1955), one variety of Anaxagorea javanica Blume (Maas & Westra, 1984, 1985), and several species of Uvariopsis (Fries, 1959). Other examples ofdimerous flowers occur in the family (cf. Fries, 1959), and in fact both trimerous and dimerous flowers occur within the magnoliid families Winteraceae, Lauraceae, Ranuncu- laceae, Berberidaceae, and Papaveraceae as well (Kubitzki, 1987).

The homogeneity of Disepalum sens. strict, and the species that have been assigned to Enicosanthellum with respect to morphology of the leaves, inflorescences, stamens, pollen, carpels, fruits, and seed argues strongly, I believe, for inclusion of all species in a single genus. The group thus defined consists of species of conservative vegetative and fruit morphologies that have evolved a spectrum of floral specializations.

Phyletic Considerations

A cladogram for the species of Disepalum was constructed using the manual quantitative parsimony algorithm described by Brooks et al. (1984) and is shown in Figure 3. Polyalthia, because of its status as a relatively "generalized" genus in the family, to which Disepalum has been allied (Sinclair, 1955), was initially used as the outgroup for purposes of establishing polarities for seven characters chosen as being unlikely, for developmental reasons, to exhibit high homoplasy. In the case of character "D" polarity was determined by its congruence with the polarities of the states of character "E"; the character therefore cannot be viewed as completely independent.

The genus Polyalthia is widely viewed as paraphyletic, and the similarities between it and Disepalum are largely superficial ones; it may thus not be the most appropriate outgroup with which to compare Disepalum. A solution to this prob- lem is to repeat the analysis using an alternative outgroup. The Cymbopetalum Tribe (sensu Walker, 1971) minus Disepalum was chosen for this purpose, because of the relationship proposed on the basis of pollen morphology. In this case, the


pet plag pul plat cor acu Ion aci ano

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FIG. 3. Cladogram for the species of Disepalum. The taxa are represented by abbreviated specific epithets, and the synapomorphies by the letters of the characters: A. Sepals three (0) versus two (1). B. Petals free (0) versus at least some petals connate (1). C. Petals free from receptacle (0) versus adnate to the receptacle (1). D. Petals six (0) versus 4-10 and variable (1) versus four and constant (2). E. Corolla lobes ovate (0) versus linear (1) versus hnear with thickened apices (2). F. Monocarps sessile on receptacle or stipitate (0) versus sessile on carpophores (1). G. Pollen grains solitary (0) versus in octads (1). Note: The character state of character D for Disepalum acuminatissimum cannot be determined from the material available.

polari t ies for all charac ters except " G ' " (pollen in octads) r e m a i n the same, and the c l adogram thus has the ident ical conf igura t ion as tha t cons t ruc ted using Polyal- thia as the ou tgroup . For charac te r " G , " oc tads are also f o u n d in the outgroup, and thus canno t be cons idered to be an a u t a p o m o r p h y defining Disepalum. O n the o ther hand, ovu le n u m b e r (4 -32 in the ou tgroup , (1) 2 (3) in Disepalum) now prov ides a new a u t a p o m o r p h y for Disepalum. Addi t iona l genera with possible re lat ionships to Disepalum can be subs t i tu ted in the same way, p rov id ing addi- t ional tests o f the hypo thes i s o f re la t ionships shown in Figure 3; w i thou t new sources o f data, however , the conf igura t ion o f the c l adogram is unl ikely to change.

A sho r t com i ng o f this analysis is tha t it leaves two unreso lved t r i cho tomies in the c ladogram, D. plagioneurum, D. petelotii, and D. pulchrum in one, and D. aciculare, D. longipes, and D. anomalum in the other. The differences a m o n g species in each o f the two t r iads are largely quant i t a t ive ones to which it is difficult to assign charac te r polarit ies, and which are highly likely to be homoplas ious . I thus, for the present , v iew the nine possible comple te ly reso lved c ladograms as equally likely a nd phylogenet ica l ly un in fo rma t ive . N e w da ta m a y change this assessment .


The cladistic analysis supports the generic circumscription proposed here. The species that have previously been included in Enicosanthellum share no derived features (despite their similarities on phenetic grounds), but are united with Di- sepalum sens. strict, on the basis of two synapomorphies. The unique progression of floral specialization found in this group suggests refinement of a functional adaptation. Many species of Annonaceae have floral morphologies adapted to very specific pollinators (Gottsberger, 1988; Murray, 1988); such may be the case here, but supporting field observations are lacking.

Several ecological and geographical patterns also emerge from the cladogram. The species occurring exclusively in montane forests, D. petelotii, D. plagioneurum, D. pulchrum, and D. platypetalum, are the more primitive ones; the more advanced species, with the exception of D. longipes, occur in lowland habitats, although D. coronatum sometimes reaches elevations of 900 meters. Disepalum coronatum and D. anomalum have become further specialized in lowland kerangas forests on the white-sand sites.

The more primitive species are also those occurring on the mainland, with the more advanced species, again with the exception of D. longipes, occurring on the islands of Sumatra and Borneo. This pattern suggests an early diversification of the genus in montane regions on the mainland of Asia, followed by a secondary dispersal and radiation of more specialized lowland species in the Malesian region. The occurrence of the specialized species D. longipes in montane habitats on the Malay Peninsula can be interpreted as either a re-diversification of the genus into a montane habitat or as persistence in the ancestral habitat. The former is the more parsimonious explanation, and is supported by Corner's (1978) observations on the lowland affinities of the summit flora of Gunung Panti, where D. longipes is known to be common.

Walker (1971, 1972) postulated that the Annonaceae originated in west Gon- dwanaland. The pattern of dispersal of the family into the Asian tropics and its subsequent diversification is, however, unclear. KeBler (1988) has recently suggested that the Asian genus Orophea Blume, the majority of whose species occur west of Wallace's Line, originated in Laurasia rather than in Gondwanaland, suggesting a dispersal of Annonaceae into Asia via the Northern Hemisphere continent. Disepalum exhibits this pattern even more strongly, with all members of the genus occurring west of Wallace's Line, four of them on the Asian mainland. The remainder of the Asian genera have not been explored in sufficient detail to reveal a recurring pattern.

The classification of the Annonaceae is currently in a state of flux, as the older systems based primarily upon perianth morphology are amended using a wider array of characters. Disepalum, as circumscribed here, underscores the shortcom- ings of the older systems, where taxa with free, homomorphic petals (the "Uno- neae" of these classifications) are classed apart from those with more specialized corollas. With this shifting of criteria for classification it is difficult to establish the affinity of Disepalum to other genera. The available evidence, especially from pollen, fruit, and seed morphology, does not support an alliance to Polyalthia. Similarly, the support for inclusion of Disepalum in the Cymbopetalum Tribe derives solely from pollen morphology (Walker, 1971), with all other evidence arguing against it. In this context it is interesting to note that the known chromosome numbers for Polyalthia (Okada & Ueda, 1984), and for Cymbopetalum Benth. and Porcelia Ruiz & Pav6n, both members of the Cymbopetalum Tribe (Morawetz, 1984, 1986), are based upon nine; the only available count for Di- sepalum is 2n = 16 for D. platypetalum (Okada, 1984).

While pollen, fruits, and seeds do not suggest a relationship of Disepalum to


Polyalthia, they do offer starting points for future phylogenetic comparisons of Disepalum to other genera. TEM studies of pollen wall ultrastructure were found useful by Le Thomas (1980, 198 l) in distinguishing superficially similar pollen types from one another in African Annonaceae; the pollen of Disepalum, because of its apparent similarity to that of the Cymbopetalum Tribe, should be examined in a similar way. Other genera with carpophores should be compared more closely: Neostenanthera and Enantia have already been mentioned, and P. Maas and collaborators (pets. comm.) report the presence of an "apical articulation" of the monocarp stalk in the genera Cananga Hook. f. & Thorns., Cleistopholis Engler, and Lettowianthus Diels, and in some species of Guatteria Ruiz & Pavrn. In Guatteria this articulation appears to be a drying artifact, not consistently present even within species, and the stalk is also articulated at the base, suggesting that the stalk is carpellary in origin and thus not homologous with the carpophore of Disepalum (pers. obs.). In the remaining genera, however, the homology of the stipe has not been established. The seeds in Disepalum are superficially similar to those found in Uvaria and Goniothalamus (Blume) Hook. f. & Thorns.; detailed comparisons of the seeds of these and other genera for characters such as endosperm configuration, histology of the testa, affd exostome morphology should provide additional data useful in establishing the position of Disepalum more firmly within the Annonaceae.

Taxonomy DISEPALUM Hook. f.

Disepalum Hook. f., Trans. Linn. Soc. London 23: 156. 1860. TYPE: Disepalum anomalum Hook. f.

Enicosanthellum Ban, Bot. Zurn. (Moscow & Leningrad) 60:808-812.1975. TYPE: Enicosanthellum petelotii (Merr.) Ban (=-Polyalthia petelotii Merr.).

Trees or shrubs with simple hairs. Leaves distichous, chartaceous to coriaceous, less commonly membranous, blade often decurrent on base, midvein prominent and usually keeled abaxially. Inflorescence terminal, less frequently leaf-opposed, 1-3-flowered, the flowers borne on slender pendent ebracteolate pedicels, the leaves opposing the pedicels often much reduced in size and the inflorescences then appearing bracteate. Sepals 5-21 mm long, foliaceous, chartaceous to subcoriaceous, slightly to deeply concave. Petals either large (up to 35 mm long) and free to the base, or smaller, ovate or strap-shaped, and united into a broad tube that is adnate to the torus for most of its length. Stamens oblong to clavate, the connectives depressed-globose and often overhanging the tops of the anther thecae. Carpels numerous, the style usually markedly narrower than the ovary; ovules usually 2, less frequently l or 3, attached laterally; style oblong, usually hairy at apex. Torus usually much wider than high, often slightly depressed in the center, often increasing markedly in size in fruit. Fruit an aggregate of sessile thinly fleshy monocarps, each subtended by a separate long stalk derived from the torus (carpophore), and separated from it by a distinct articulation. Monocarps two-seeded, less frequently one-seeded or rarely three-seeded in D. pulchrum, the monocarp wall separating readily from the seed in the dried condition. Seeds ellipsoid or flattened ellipsoid, surface in contact with adjacent seed flattened, chestnut-brown, glabrous, shiny, the rim of the exostome papery, light colored, and slightly re- cessed; seed coat with numerous internal plates extending between the ruminations of the endosperm.

Nine species, distributed from southern China south to Sumatra and Borneo, at elevations from sea level up to 2500 meters.

364 aRITTONIA [VOL. 41

K e y to the spec ies o f D i s e p a l u m

1 Sepals 3; all petals free from one another at base; neither sepals nor petals persistent in fruit. 2 Carpels 18-35; carpophores 0.7-1.3 cm long; petals 13-26 mm long .......................................................

........................................................................................................................................................ 1. D. pe te lo t i i (Merrill) D, M. Johnson 2 Carpels 60-200; carpophores at least 2 cm long; petals 21--44 mm long.

3 Monocarps broadly cuneate to rounded at base; leaf yellow-green beneath; China, Viet- nam ...................................................................................................................... 2. D. p l a g i o n e u r u m (Diels) D. M. Johnson

3 Monocarps attenuate and narrowly cuneate at base; leaves sometimes paler beneath, but green; Malay Peninsula ................................................................................ 3. D. p u l c h r u m (King) J. Sinclair

1 Sepals 2; at least some of the petals connate for half their length and adnate to the sides of the torus; both sepals and petals usually persistent in fruit.

4 Corolla lobes as wide or wider than long; monocarps 11-23 mm long ................................................... ............................................................................................................................................................................................. 4. D. p l a t y p e t a l u m Merr.

4 Corolla lobes longer than wide; monocarps 6-13 mm long. 5 Corolla lobes 4--9, 7-12.5 mm long, tapering to an apex of uniform thickness.

6 Acumen of leaf4-10 mm long; corolla glabrous, or with sparse, pale erect or appressed hairs; Borneo ............................................................................................................................................... 5. D. coronaturn Becc.

6 Acumen of leaf 15-18 mm long; corolla with abundant rusty appressed hairs; Sumatra ..................................................................................................................................... 6. D. a c u m i n a t i s s i m u m Boerl. & Koord.

5 Corolla lobes 4, 3-8 mm long, wider and slightly thickened toward apex. 7 Corolla lobes acute at apex, with a maximum width of 0.5-0.6 mm; styles with

crisped hairs at apices ............................................................................................. 7. D. acieulare D. M. Johnson 7 Corolla lobes blunt at apex, with a maximum width of 1-2.3 mm; styles with straight,

rigid hairs at the apices. 8 Expanded apex of corolla lobe longer than wide; stamens 1-1.5 mm long; Malay

Peninsula, Sumatra ............................................................................................................................... 8. D. longipes King 8 Expanded apex of corolla lobe as wide as or wider than long; stamens 1.5-2.1 mm

long; Borneo ................................................................................................................................. 9. D. a n o m a l u m Hook. f.

1. D i s e p a l u m pe te lo t i i (Mer r . ) D. M. J o h n s o n , c o m b . nov . (Figs. 1A, 2A)

Polya l th ia pete lot i i Merrill, Univ. Calif. Publ. Bot. 13: 131-132. 1926. E n i c o s a n t h e l l u m pete lo t i i (Merr.) Ban, Bot. Zurn. (Moscow & Leningrad) 60: 812. 1975. TYPe: INDOCHINA. T o ~ [VmTNAM]: Chapa, P~telot 1780 (HOLOTVa~E: UC, fragment and photograph at A!; ISOTVVES: At, NY!).

S h r u b o r sma l l t ree 2 - 5 m tall . Lea fy p o r t i o n o f twigs 1 .3-3 m m th ick , g labra te , r e d d i s h - b r o w n to d a r k gray, l o n g i t u d i n a l l y w r i n k l e d . L a m i n a o f la rger l e a v e s 14 .6 - 17.5 c m long, 3 .3 -4 .6 c m wide , cha r t aceous , o b l a n c e o l a t e to n a r r o w l y e l l ip t i c o r ob long , base cunea te a n d s h o r t d e c u r r e n t on pe t io le , apex a c u m i n a t e , t he a c u m e n 9 - 2 0 m m long, g l ab rous adax i a l l y , i n i t i a l ly g o l d e n to r u s ty - s e r i c e ous a ba x i a l l y , a t l ength g labra te or w i th s ca t t e r ed a p p r e s s e d ha i rs ; m i d r i b i m p r e s s e d adax iaUy, p r o m i n e n t a n d kee led abax i a l l y , s e c o n d a r y v e i n s 8 - 1 3 pe r s ide, these a n d h ighe r o r d e r ve ins s l ight ly r a i s ed on b o t h surfaces. Pe t i o l e 4 - 7 m m long, 1 .2 -1 .6 m m wide , pubescen t . In f lo rescence t e r m i n a l , l - f l o w e r e d ; pe d i c e l 1-1 .7 c m long, 1.3 m m t h i c k at m i d p o i n t , f e r r u g i n o u s - p u b e s c e n t . Sepa l s 3, 10-11 m m long, 8 -9 .5 m m wide , cha r t aceous , ova t e , acu te at apex , a p p r e s s e d - p u b e s c e n t on b o t h surfaces , e v e n t u a l l y g lab ra te abax ia l ly . Pe ta l s free, in 2 w h o r l s o f 3, co r i aceous , o b l o n g o r ob l anceo l a t e , acu te at apex , g l ab rous to s p a r s e l y p u b e r u l e n t adax i a l l y , spa r se ly se r i ceous abax ia l ly ; o u t e r pe ta l s 14-26 m m long, 6 .5 -11 m m wide; i n n e r pe ta l s 13 -23 m m long, 7-11 m m wide . S t a m e n s 2 . 2 - 2 . 7 m m long; c o n n e c t i v e s de - p re s sed -g lobose , s h o r t - p u b e s c e n t . Ca rpe l s 18-35 ; o v a r y 0 . 5 - 1 . 2 m m long, ap - p r e s sed pubescen t ; s ty le 1 .2-1 .5 m m long, h i s p i d u l o u s a t apex; o v u l e s 1 o r 2. T o r u s 4 - 1 0 m m in d i a m . Ped ice l in f ru i t ing s tage 1 .3-2 .2 c m long, 1 -2 .3 m m t h i c k a t m i d p o i n t , f e r r u g i n o u s - p u b e s c e n t . T o r u s in f ru i t ing stage 4 . 5 - 6 m m in d i a m , p e r i a n t h no t pe r s i s t en t ; c a r p o p h o r e s 0 . 7 - 1 . 3 c m long, 0 . 9 - 2 m m t h i c k a t m i d p o i n t , ob l i que ly c l ava t e to t runca te , 1 .5 -2 .3 m m t h i c k a t apex . M o n o c a r p s 4 - 1 5 , c o n c o l o r o u s wi th c a r p o p h o r e s o r da rke r , 1 3 - 1 4 m m long, 7 - 9 m m wide , e l l i p s o i d to ob long -e l l i p so id , apex r o u n d e d , b a s e r o u n d e d o r s l ight ly a t t enua t e ,


P E O P L E ' S R E P U B L I C OF C H I N A

NAM

96 ~ 1 2 o +

200km

s% ; IA

�9 D. peteloti i

* D. p l a g i o n e u r u m

�9 D. pu l ch rum

FIo. 4. Distributions of Disepalum petelotii, D. plagioneurum, and D. pulchrum in the People's Republic of China, Vietnam, Laos, and Malaysia.

glabrate; monocarp wall ca 0.1 mm thick. Seeds 1 or 2, 10-12.5 mm long, 6-8 mm wide, flattened-ellipsoid.

Phenology and distribution: Flowering collections of this species are all from April, and the fruiting collections are from August and September. This species is known from central Laos, northern Vietnam, and southern Yunnan, People's Republic of China, where it occurs in montane evergreen forests at elevations of 1200 to 1900 meters (Fig. 4).

Color notes: Petals greenish white, with brown patch at base (Kerr 21162). Monocarps black (Henry 10886).

366 BRITTONIn [VOL. 41

Additional specimens examined: PEOPLE'S REPUBLIC OF CHINA. Ytn~AN: Si-chour-hsien [Si- chou], Faa-dou [Far-dou], Feng 11885 (A); Mar-li-po, Sze-tai-po, Loa-chun-shan, Feng 13818 (A); Mengtse, Henry 10886 (A, NY). VIETNAM: Chapa, P~telot 5471 (A, NY); Prov. Lao Kay, Ta Yang Binh, P~telot 7638 (A). LAOS: Pu Muten, Chieng Kwang [Xiangkhoang], Kerr 21162 (K); Pu [Phou] Bia, Kerr 21783 (K).

Disepalum petelotii differs f rom D. plagioneurum in its smaller petals, lower carpel number, and shorter carpophores of the fruits. Where the two occur sym- patrically in northern Vietnam, D. petelotii occurs at higher elevations.

A synonym listed for this species by Tsiang and Li (1979) is Uvaria oblanceolata W. T. Wang (Acta Phytotax. Sin. 6(2): 197-198, 1957). I have been unable to see the type o f this name, but there are elements o f the protologue (e.g., six ovules per carpel) that make it unlikely to represent this species; one of the paratypes, Feng 11885, is, however, D. petelotii. The combina t ion Disepalum petelotii was made inadvertently by Klucking (1986), who failed to cite the basionym; the combinat ion is thus formally proposed here.

2. Disepalum plagioneurum (Diels) D. M. Johnson, comb. nov. (Figs. 1B, 2B)

Polyalthia plagioneura Diels, Notizbl.Bot. Gart. Berlin-Dahlem 10: 886-887. 1930. Enicosanthel- lure plagioneurum (Diels) Ban, Bot. Zurn. (Moscow & Leningrad) 60:812.1975. TYPE: PEOPLE'S REPUBLIC OF CHINA. GUANGXI: Yao-shan, 1600 m, 27 Apr 1929, Sin 8145 (HOLOTVPE: B, photograph and fragment at A!; XSOTYPE: IBSC!).

Uvaria petelotii Exell, J. Bot. 70: 104. 1932. TYPE: INDOCHINA: Tonkin, massif of Tam Dao, about 900 m, May, P~telot 3964 (HOLOTYPE: BM!, fragment at A!).

Shrub or tree 3-18 m tall with a DBH of at least 2-3 cm. Leafy port ion o f twigs 1.5-5.2 m m thick, sparsely pubescent at first, soon glabrate, brown to dark gray or purplish-gray, longitudinally wrinkled. Lamina o f larger leaves 10.6-18 cm long, 3-7.1 cm wide, chartaceous to subcoriaceous, elliptic, less commonly oblong, oblanceolate, or lanceolate, base cuneate and decurrent, apex acute or short- acuminate, the acumen 3-9 m m long, glabrous adaxially, at first densely rusty sericeous abaxially, soon sparsely appressed-pubescent; midrib slightly impressed adaxially, raised and keeled abaxially; secondary veins 8-12 per side, these and higher-order veins p rominen t on both surfaces. Petiole 6-10 m m long, 1.5-2.3 m m wide, pubescent to glabrate. Inflorescence terminal, 1-flowered; pedicel 2 .2- 4.3 cm long, 1.5-2 m m thick at midpoint , rusty-pubescent. Sepals 3, 14-18 m m long, 13-17 m m wide, chartaceous, broadly ovate, obtuse to apiculate at apex, pubescent to sparsely pubescent. Petals free, in 2 whorls o f 3, chartaceous, oblanceolate, acute at apex, adaxially with dense erect hairs and petal surface b u m p y creating an appearance o f stellate hairs, uniformly appressed-pubescent abaxially; outer petals 22-32 m m long, 9-19 m m wide; inner petals 23-34 m m long, 12- 24 m m wide. Stamens 2.2-2.6 m m long; connectives cushion-shaped, puberulent. Carpels 60-200; ovary ca 1 m m long, rusty appressed-pubescent; style 1.4-1.8 m m long, pubescent at apex; ovules 1 or 2. Torus 7-9 m m in diam. Pedicel in fruiting stage 2.8-5.4 cm long, 1.7-4 m m thick at midpoint , pubescent. Torus in fruiting stage 11-21 m m in diam; perianth not persistent; carpophores 3-4.7 cm long, 1.5-1.7 m m thick at midpoint , t runcate and 3-5 m m thick at apex. Mono- carps (3) 10-98, darker than carpophores and slightly glaucous, 11.5-16 m m long and 8-11 m m wide, ellipsoid, apex rounded, base broadly cuneate to rounded, glabrate; monocarp wall 0.5-0.6 m m thick. Seeds 1 or 2, 11-11.5 m m long, 8 m m wide, ellipsoid.

Phenology and distribution: Flowering collections of D. plagioneurum have been made from March through May, while fruiting specimens have been collected in June, October, December, and March. The species occurs in lower montane forests at elevations o f 600 to 1600 meters, at scattered localities in the southern

1989] JOHNSON" DISEPALUM 367

People 's Republ ic o f China, including the island o f Hainan , and in nor thern Vie tnam (Fig. 4).

Color notes: Flower color is given as pale green to yellow, fruit color as black.

Additional specimens examined: PEOPLE'S REPUBLIC OF CHINA. GUANGDONG: Hainan, Yai- chow, How 70331 (A, NY); Hainan, Po-ting, How 72863 (A); Yeung Ling Shah, Ngai [Yai] District, Lau 101 (A, K, MICH, NY); Chim Fung Mt., near Fong Ngan Po Village, Kan-en District, Lau 5652 (A); Bak Sa [Pai-sha], Lau 25740 (A); Wentongshan, Liang 60660 (NY); Hainan, Liang 63386 (A, NY); Hainan, Po-ting Pref., Suila Mountain, Hsin-an Village, Suila Mountain Climbing Party 3193 (PE); Ting-wu Shan, Ting & Shih 1086. GUANGXI: Yao Shan, Wang 40300 (A). VIETNAM: Tam Dao, prov. Vinh Jen, Eberhardt 4964 (A); prov. de Quang Nam [Annam], Poilane 29564 (A, L); prov. du Vinh Jen, massif du Tam Dao, P~telot 3954 (A, NY), Chapa, 6362 (A, MICH, NY).

The larger flowers with more numerous carpels, along with longer carpophores (and more numerous fruits), set this species apart f rom D. petelot iL the other northern species o f Disepa lum. As far as is known, the two species over lap in range only in nor thern Vietnam.

3. DISEPALUM PULCHRUM (King) J. Sinclair (Figs. 1 C, 2C)

Polyalthia pulchra King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61: 55. [13 Jun] 1892. Disepalum pulchrum (King) J. Sinclair, Gard. Bull. Straits Settlem., Ser. 3, 14: 333. 1955. Enicosanthellum pulchrum (King) van Heusden, Proc. Kon. Ned. Akad. Wetensch., Ser. C, 91(3): 248. 1988. TYPE: MALAYSIA: Perak, Weld's Resl, Scortechini 824 (HOLOTYPE: K, not seen; ISOTYPE: CAL, photograph!).

Polyalthia pulchra var. angustifolia King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61: 55. [13 Jun] 1892. Disepalum pulchrum vat. angustifolium (King) J. Sinclair, Gard. Bull. Straits Settlem., Ser. 3, 14: 334. 1955. TYPE: MALAYSIA: Perak, Gunung Bubu, 5000 It, Mar 1890, Wray 3867 (HOLOTYPE: K, not seen; ISOTYPES: BM!, CAL, photograph!, SING!).

Tree 3-14 m tall with a D B H of 15-30 cm. Leafy port ions o f twigs 1.3-4.6 m m thick, appressed-pubescent to glabrescent, reddish to grayish-brown, longitudinally striated to wrinkled, occasionally with oblique longitudinal ridges decurrent f rom petiole base. L a m i n a o f larger leaves 12.7-21.5 cm long, 4.5-8.1 cm wide, chartaceous, obova te to broadly oblanceolate, occasionally elliptic to oblong, base cuneate and short-decurrent , apex shor t -acuminate , with a blunt acumen 4-11 m m long, glabrous adaxially, appressed-pubescent abaxially, eventual ly very sparsely pubescent to glabrescent; mid r ib slightly impressed adaxially, raised and keeled abaxially; secondary veins 8-12 (15) per side, these and higher-order veins slightly raised on bo th surfaces. Petiole 4 -12 m m long, 1.5-2.1 m m wide, pubescent. Inflorescence terminal , 1-flowered; pedicel 3.6-5.3 m m long, 1.2-1.7 m m thick at midpoint , appressed-pubescent . Sepals 3, 10o21 m m long, 7.5-15 m m wide, chartaceous, ovate, acute at apex, appressed pubescent on both surfaces. Petals free, in 2 whorls o f 3, subcoriaceous, oblanceolate or pandur i fo rm, acute to cuspidate at apex, erect-pubescent to glabrous adaxially, appressed-pubescent abaxially, 21-44 m m long, 8-25 m m wide. Stamens 2.8-3.8 m m long; connect ive cushion-shaped, erect-pubescent. Carpels 1200200; ovaries 2.1-2.9 m m long, densely pubescent; styles 1.2-4.5 m m long, slightly setulose at apex; ovules 2, rarely 3. Torus 7-8.5 m m in diam. Pedicel in fruiting stage 5.3-6.7 cm long, 2 .5- 4.3 m m thick at midpoint , sparsely appressed-pubescent . Torus in fruiting stage 11-25 m m in diam; per ianth not persistent; carpophores 3.1-4.4 cm long, 1.3- 2.1 m m thick at midpoint , clavate and 2.2-3.5 m m thick at apex. Monocarps 13- 102, concolorous with carpophores, 14-21 m m long and 7-10 m m wide, ellipsoid, apex rounded or apiculate, base at tenuate, glabrescent; m o n o c a r p wall 0.1-0.2 m m thick. Seeds 2, less c o m m o n l y 1 or 3, 10-13 m m long, 7.5-8.5 m m wide, flattened-ellipsoid.

Local names: m e m p i s a n g , bunga pisang, t a m p i n g gunong.


Phenology and distribution: Disepa lum p u l c h r u m is in flower and fruit throughout the year. This species is fairly c o m m o n in the highlands o f the nor thern Malay Peninsula, at elevations o f 1200 to 1900 meters (Fig. 4).

Color and fragrance notes: Flower color as repor ted on collection labels is variable. The calyx is always green, but the petals are described as greenish-yellow, yellow, c reamy yellow, cream, peach-colored and brownish-orange outside and brownish-purple inside. The petals, at least the inner ones, are repor ted to have a chevron-shaped reddish-purple patch on the inner surface toward the base. Anther connectives are repor ted to be white or yellow, and styles green or brownish-pink. It is probable that the var ia t ion in flower color reflects different stages in deve lopment , especially toward anthesis, but field studies are required to de- t e rmine this. Floral fragrance is r emarked upon by several collectors: the label o f H o l t t u m 31316 describes it as "delicate, like a ripe apple ," while S t one 14427 called it "tart-sweet, tending to a bit sickly." The impress ion of a fruity smell is given in all cases. Van Balgooy 2645 ment ions the presence of beetles in the flowers.

The monocarps are described as var ious shades o f purple, and are borne on pinkish-red, cr imson, or purpl ish carpophores.

Additional specimens examined: MALAYSIA. KELANTAN: Gunong Rabong, Soepadmo & Mahmud 1105 (KEP, SING). PAHANG: Cameron Highlands, track to Robinson's Falls, near Tanah Rata, van Balgooy 2645 (NY); Cameron Highlands, Sg. Uruil, below Golf Course, Burkill HMB.813 (A, L, SING); Pine Tree Hill near Fraser Hill, upon the Selangor border, Burkill & Holttum SFN 8533 (SING); Mentigi FR, Cameron Highlands, Chan FRI 16995 (KEP); Path 1, Tanah Rata, Cameron Highlands, Chew 236 (A, C, NY, SING); Cameron Highlands, Bukit Siku, 4~ 101~ Chew 824 (A, L, SING); Cameron Highlands, Bukit Mentigi, Chew 857 (A, K, SING, UC); Cameron Highlands, foot of Gunong Berembun, Chew 1249 (A, K, L, SING, UC); Ginting [Genting] Highlands near the Selangor border, Chin 1207 (KLU); Mentigi Nursery, Cameron Highlands, Durant 28734 (KEP); near Tanah Rata, Cameron's Highlands, Henderson SFN 17736 (SING); Cameron's Highlands, Henderson SFN23288 (BM, KEP, NY, SING); Sungai Terla, Holttum SFN31316 (A, K, KEP, SING); S. Terolak, Ulu Gelom, Jaamat 27574 (KEP, SING); Ulu Sg. Lemoi, Jaamat 28116 (KEP, SING); Cameron Highlands, Tanah Rata, Ja'amat & San 36179 (KEP); Cameron Highlands, path to G. Jasar, Kochummen FRI 19378 (KEP, L); Cameron Highlands, Gunong Beremban, Kochummen KEP 99971 (A, KEP); Cameron Highlands, Sample Plot I, Ng FRI 5882 (A, L); Cameron Highlands, stream to Tanah Rata, Poore 1014 (KLU); Teku River, Gunong Tahan, Ridley 16012 (BM, SING); Cameron Highlands, path off mile 388 to Parit Falls, Sungei Jasar, Tana Rata, Sinclair 9930 (A, L, SING); Cameron Highlands, Robinson's Falls, Stone 14427 (A, KEP, KLU); Gunung Beremban, Stone 14455 (A, KEP, KLU); Genting Highlands road below Ulu Kali summit, montane forest opposite Sri Genting Villa, Stone 15371 (KLU); Gunong Brinchang, Cameron Highlands, Student 8086 (KLU); Cameron Highlands, Wyatt-Smith 63655 (KEP); Mentigi Plantations, Cameron Highlands, Wyatt-Smith 71922 (A, K, KEP, L, SING); Three Hill, Fraser Hill, Collector unknown 22437 (KEP). PAHAN~/SELANGOR: about 1 mi below the summit of Ulu Kali, Anderson 2949 (MO); Ginting [Genting] Highlands near Hotel, Kochummen FRI 16735 (A, K, KEP); Summit Plateau, G. Ulu Kah [Kali?], Whitmore FRI 12211 (K, KEP, L, SING). PERAK: Gunong Kerbau, 1913, H. C. Robinson s.n. (K, SING). SELANC, OR: Gunong Ulu Kali near wireless station, Genting [Ginting] Highlands, Shah & Ali MS 2939 (HBG, KEP, SING). STATE UNKNOWN: G. Mengkuang, Wyatt-Smith KEP 93114 (A, K, KEP).

This species is mos t s imilar to D. p lag ioneurum, but differs in the shape o f the monocarps , and in the concolorous rather than discolorous leaves. There is also a tendency for the petals o l D . pu lchrum to be cuspidate at the apices ra ther than acute as in D. p lagioneurum, but this is not a constant feature.

The narrow-leaved var ie ty o f this species recognized by bo th King and Sinclair is reduced to a synonym of the species here. Recent collecting has produced a good series of specimens o f D. pulchrum, some o f which fill the gaps in leaf and petal widths between D. p u l c h r u m var. p u l c h r u m and D. p u l c h r u m vat . angusti- f o l i um. The type of D. p u l c h r u m var. angus t i fo l ium thus appears to represent an ex t reme in a con t inuum o f variat ion.


VIALAY

oNIN ULA

�9 D. p l a t y p e t a l u m

D. acum ina t i s s imum

o D. Iong ipes

110~ 200km

Cb

D. c o r o n a t u m

�9 D. a c i c u l a r e

�9 D. a n o m a l u m

FIG. 5. Distributions of Disepalum platypetalum, D. acuminatissimum, D. longipes, D. coronatum, D. aciculare, and D. anomalurn on the Malay Peninsula, Sumatra, and Borneo.

4. DISEPALUM PLATYPETALUM Merrill (Figs. 1 D, E, 2D)

Disepalum platypetalum Merrill, Contr. Arnold Arbor. 8: 58. I934. TYPE: SUMATRA, Tapianoeli, road from E Coast to Tapianoeli, NW side of Toba Lake, near Peso Peso, in primary forests, alt. 1200-1500 m, 16 Feb 1932, Bangham & Bangham 1077 (LECTOTYPE, designated here: A!; ISOLECTOTYPE: NY!).

Tree or weak bent shrub, 4.5-9 m tall. Leafy port ion of twigs 0.9-3.3 m m thick, initially glabrous, or with sparse very fine pubescence and soon glabrate, reddish- or purplish-brown to dark gray, longitudinally wrinkled. Lamina o f larger leaves 10.5-18 cm long, 3-5.8 cm wide, chartaceous to subcoriaceous, oblanceolate or rarely oblong, base cuneate and short-decurrent on petiole, apex cuspidate or short-acuminate, with a blunt cusp or acumen 4-15 m m long, glabrous adaxially, initially golden-sericeous or with fine appressed hairs abaxially but soon glabrate; midrib impressed adaxially, p rominent and sharply keeled abaxially, secondary veins 11-16 per side, these and higher-order veins raised on both surfaces. Petiole 7-15 m m long, 0 .9-15 m m thick, glabrous. Inflorescence terminal or less frequently leaf-opposed, 1-flowered; pedicel 3.3-8.9 cm long, 0.5-1 m m thick at midpoint , with scattered appressed hairs, or glabrous. Sepals 2, 9-12 m m long, ca 9-9.5 m m wide, subcoriaceous, orbicular or broadly ovate, rounded at apex, densely pubescent adaxially, sparsely pubescent abaxially. Petals connate or sometimes both free and connate in the same flower, coriaceous; tube 2.5-4.2 m m long, partially adnate to torus at base, glabrous; corolla lobes 5-9, 2-2.8 m m long, 4.2-5.5 m m wide at widest point, broadly triangular, obtuse at apex, glabrous. Stamens ca 2 m m long; connectives depressed-globose, short-pubescent. Carpels 70-200; ovary 1-1.4 m m long, appressed-pubescent; style 1.4-2 m m long, setulose at apex; ovules 2. Torus 8 m m in diam. Pedicel in fruiting stage 4.6-8.4 cm long,


1.3-1.7 m m thick at midpoin t , glabrate. Torus in fruiting stage 1.2-2.1 cm in d iam, often with calyx and /o r corolla persistent; carpophores 2.2--4.5 cm long, 0.6-1.7 m m thick at midpoin t , thickened, obliquely clavate, and 1.2-3.7 m m thick at apex. Monocarps 22-69, concolorous with carpophores , 11-23 m m long and 5.5-11 m m wide, ellipsoid, apex rounded to mucronate , base rounded or slightly attenuate, glabrate; m onoca rp wall 0.2 m m thick. Seeds 1-2, 8-11 m m long, 5.5-9 m m wide, irregularly ellipsoid.

Local names: andor p a k p a k saloeng, andor bosi. Phenology and distribution: Flowering collections o f D i s e p a l u m p l a t y p e t a l u m

are f rom February, March, and June, while fruits have been collected in February, March, June, and Oc tobe r -December . This species is restricted to m o n t a n e rain- forest at 600 to 2500 m in the vicinity o f T o b a Lake, nor thwestern Sumat ra (Fig. 5).

Color notes: The photograph published in Ho t t a (1989) shows the sepals o l D . p l a t y p e t a l u m to be green, the petals to be brownish-cream, and the anther connectives to be pale brown; one collector describes the flowers as having "bracts [sepals?] deep m a r o o n ? ' The fruit is reported to be purplish red or scarlet.

Additional specimens examined: SUMATRA: Road NW of Toba Lake, Bangham & Bangham 1232 (A, lectoparatype); E coast, lower slopes of Dolok Si Manoek-manoek, Asahan (NW from Taloen na Oeli, Toba), Boeea 10239 (A, MICH, NY); res. Tapianoeli, Dolok Ri da Bolak, Toba (N of Hoeta Djoeloe, on the trail from Loemban Loboe to Taloen na Oeli), Boeea 11251 (A, MICH, NY); Atjeh, lower elevation of Gunong Kemiri, Iwatsuki et al. S.808 (KYO); Atjeh, higher elevation of Gunong Kemiri, Iwatsuki et al. S.1074 (KYO); in silvis prope Koeroeng [?], Junghuhn s.n. (L); res. Atjeh, Gajolanden, Sanger valley to bivouac 2, above Blang Kedjeren, van Steenis 9843 (A, K, NY, P, SING); Atjeh, Gunung Leuser Nature Reserves, Upper Mamas River valley, ca 15 km W of Kutacane, 3"25'N, 97"40'E, de Wilde & de Wilde-DuyJ~es 18346 (KYO), 18353 (L), 18622 (KYO, L), 19092 (L).

Like D i s e p a l u m corona tum, D. p l a t y p e t a l u m exhibits var ia t ion in corolla lobe n u m b e r on different flowers o f the same plant. The corolla in D. p l a t y p e t a l u m m a y exhibit an addi t ional aspect o f variabi l i ty in that the petals, while always adnate to the sides of the torus, can be both free and connate in the same flower.

5. DISEPALUM CORONATUM Beccari (Figs. 1F, G, 2E)

Disepalum coronatum Beccari, Nuov. Giorn. Bot. Ital. 2: 155. 1870. TYPE: BORNEO. SARAWAK: Mt. Mattan (Mt. Matang), 2500 r, [May or Jun 1866], Beccari P,B. 1722 [cited as P. B. 1732 in protologue] (HOLOIYPE: FI, not seen; ISOTYPES: A!, B!, C!, G!, K!, NY! (fragmen0, P!).

Tree or shrub 1.5-9 m tall with a D B H of 5-30 cm; bark smooth , gray to black, somet imes mott led, inner bark pale yellow or gray, sapwood white or yellowish. Leafy por t ion of twigs 0.7--4.3 m m thick, glabrous or sparsely pubescent and soon glabrate, dark reddish or yel lowish-brown, with age dark gray or gray-brown, often with frosty exfoliating ep idermal patches, longitudinally wrinkled, somet imes slightly verrucose. L a m i n a o f larger leaves 7.8-18.4 cm long, 3.1-6.3 c m wide, subcoriaceous, occasionally coriaceous or chartaceous, oblanceolate, oblanceolate- obova te or elliptic, base cuneate and shor t -decurrent on petiole, apex with an acumen 4-10 m m long, glabrous adaxially, initially sparsely appressed pubescent or glabrous abaxially, soon glabrate; midr ib slightly impressed adaxially, raised and keeled abaxially; secondary veins 6-15 per side, c o m m o n l y 9-1 1, these and higher-order veins raised or indistinct on bo th surfaces. Petiole 4-13 m m long, 0 .8-2.2 m m wide, glabrous. Inflorescence terminal , with the opposing leaves often reduced in size, 1- or 2-flowered; pedicels 1.4-4.6 c m long, 0.6-1.3 m m wide at midpoin t , glabrous or with a few appressed hairs. Sepals 2, (5) 7-1 1 m m long, 6.5-1 1 m m wide, 3-5 m m deep, coriaceous to subcoriaceous, orbicular to ovate, often boat-shaped, rounded to obtuse at apex, densely short-oubescent adaxiaUy,


glabrous or sparsely pubescent abaxially. Petals connate, coriaceous to subcoriaceous; tube (0.5) 2-6.5 (7.5) m m long, broadly funnelform, adnate to torus at base, glabrous; lobes 4-9, (5) 7-12.5 (14) m m long, 1.7-5.5 m m wide at widest point, linear, evenly tapering to an acute or obtuse apex, sparsely appressed- pubescent only on ul t imate 1-2 m m of lobes, otherwise glabrous. Stamens (1.2) 1.5-2 m m long; connectives cushionlike, erect-pubescent. Carpels ca 60-120; ovary 0.7-1.5 m m long, appressed-pubescent ; style 0.5-1.6 m m long, setulose or hispidulous at apex; ovules (1) 2 (3). Torus 5-8 m m in d iam. Pedicel in fruiting stage 2.2-5.4 cm long, 1-2.5 m m thick at midpoint , glabrate. Torus in fruiting stage 6-20 m m in diam, calyx and corolla usually persistent; carpophores (0.6) 1.9-4.0 cm long, 0.6-1.5 m m thick at midpoint , evenly to obliquely truncate and I. 1-2.2 m m thick at apex. Monocarps (8) 17-76, concolorous with carpophores, 6-11 m m long, 5.5-8 m m wide, ellipsoid, apex rounded to apiculate, base broadly cuneate to rounded, glabrate; monoca rp wall 0.2 m m thick. Seeds 2, less commonly 1, 5 .5-12 (15) m m long, 4 .5-7 m m wide, irregularly ell ipsoid to globose.

Local names: k a t o k (Sabah), pendok, karai bunga (Brunei), ka joe sangai (Kali- mantan) .

Phenology and distribution: In flower practicdlly throughout the year; no flowering collections f rom April or May. Fruit ing collections are known f rom throughout the year as well. Nei ther flowering nor fruiting appears to have a seasonal or geographic variat ion. D i s e p a l u m c o r o n a t u m occurs on peaty or sandy soils at elevations f rom sea level to 900 meters on Borneo f rom Sabah to western Sarawak, with a couple o f outlying localities in nearby Ka l iman tan (Fig. 5). This species mos t frequently occurs in heath forests on either white sand (kerangas) or sandstone mounta ins , but is somet imes found in peat swamp forests.

Color notes: F r o m the numerous collections o f this species it is possible to piece together the sequence of changes in floral color. The sepals and petals remain green, al though the petals m a y take on a reddish-brown tinge in fruit. The anther connectives are at first green or tan, but at anthesis become bright cr imson. The styles remain c ream colored throughout , but at anthesis secrete a clear sticky fluid. In mature fruits the carpophores and torus become pink, while the monoca rps are dark red or cr imson.

Representative specimens examined: BORNEO. SAaAH: Beaufort District, 1.5 km S of Weston, Beaman 8064 (MSC, NY), 9430 (MSC, NY); Beaufort District, Lupak, Buntar SAN 25827 (SING); Beaufort District, Lumat, Dewol & Karim 77889 (K); Kimanis, Evangelista 3770 (A, SING); Tawau District, Mt. Andarassy, Gibot SAN32966 (KEP, L); Damit River, Hassan 810 (NY); Weston District, Kampong Usak, Jurnatin SAN 72519 (K); Tuaran Dist., Sg. Damit F. Res., Nic[h]olson SAN 39763 (L); Keningau, Rashna area, mile 6 Nabawan, Nordin Abas. SAN 85777 (K, KEP, L); Nabawan, Mile 41/2 Jalan Bornion Camp, Nordin & Jumatin SAN 90258 (K, L); Bangawan Forest Reserve, Rahim A428 (L, SING, US); between Bonguawan and Mandahan along railway, Stone 6755 (KLU, L); Keningau, Mile 10 Rashna road, Nabawan, Sundaling SAN 83836 (K, L); Keningau Dist., Mile 5 Rashna road, Nabawan, Sundaling SAN 83862 (K). BRtn~i: Pylon 50, Badas Railway, Ashton BRUN 983 (K, L); Badas F. R., Briinig S.1064 [S.1194?] (K, L); Tutong Dist., between Danau and Tutong, van Niel 4399 (KYO, L). SARAWA~: Baram, Anderson 2082 (KEP, SING); Baram Dist., Melinau Terraces N of Gunong Benarat [Benaret], Anderson 4614 (K, L); near summit of Santubong, Anderson 8361 (K, KEP, L, SING); Miri Dist., Lambir Hills F. R., Bakar 3050 (A, K, L, SING); Telok Asam, Bako N. P., Chai & Paie S.17845 (A, L, SING); Bkt. Goram, Ulu Sg. Kapit, 7th Div., Chai S.36128 (G, K, KEP, L); 4th Division, about 30 km S of Miri, top of Bukit Lambir, Fuchs 21330 (K, L); Lambir N. P., 4th Division, George S.40407 (K, L); Mt. Matang, Garaifor Haviland 1502 (K); Mr. Singhi, Garaifor Havitand 2024 (K); NE slope of Bt. Kana, Bintulu District, 4th Division, Hirano & Hotta 1301 (KYO); border line between Bkt. Dema and Balui, Belaga, 7th Division, Lee S.39984 (K, KEP, L); Santubong, Kuching, 1 st Div., Gunong Santubong, Martin S.37678 (L); on summit of Lambir Hills N. P., Ulu Sg. Lebau, 20th mile Miri/Bintulu Road, 4th Division, Pale & Teck S.38355 (K, KEP, L); Ulu Temiai, Mujong, Hose Mts., Sibat ak Luang S.17148 (A, K, KEP, L, SING). KAU- MA~rrAN: Oeloe Kenepai, Hallier 1432 (L); Pontianak, Kp. Andjongan, Mondi 237 (K). Borneo, unspecified: Teysmann 120 (MICH), s.n. (L).

3 72 BRITTONIA [VOL. 41

This species exhibits a great degree of floral variability, both in size of the flowers and in corolla morphology. The latter principally involves changes in the number of corolla lobes, which may be as few as four or as many as nine, with all possibilities in between documented. Different numbers can occur in different flowers on the same branch, but the possible variation over an entire tree is not known. The linear shape of the lobes is, in contrast, very constant regardless of number. The shape of the corolla tube appears to be somewhat correlated with the number of lobes; flowers with four lobes often have a more narrowly funnelform tube (Fig. 1F), while in those with six or more the tube is short and virtually obscured by the androecium (Fig. 1G).

6. DISEPALUM ACUMINATISSIMUM Boerlage & Koorders

Disepalum acuminatissimum Boerlage & Koorders in Koord.-Schum., Syst. Verz. Herb. Koord. 2: 19.19 i0. TWE: SUMATRA. "Ira Inneren des Flachmoors-Hochwaldes bei Langgam, aufmeterhoch iiberschwemmtem Moorboden, um 20 m. ii.M.," 26 Mar 1891, Koorders 157778 (nOLOa'VPE: BO!, fragment at B!).

Leafy portion of twigs 1.2-1.6 mm thick, dark reddish to yellowish-brown, nearly smooth. Lamina of larger leaves 14.4-14.7 cm long, 3.9-5 cm wide, oblanceolate, membranous, base cuneate, decurrent, apex long-acuminate, the acumen 15-18 mm long, glabrous adaxially, sparsely pubescent abaxially; midrib slightly impressed adaxially, raised and keeled abaxiaUy; secondary veins 8-10 per side, these and higher-order veins slightly raised on both surfaces. Petiole 7- l 1 mm long, 0.8-1.3 mm wide, glabrous. Pedicel up to 6.3 cm long, 0.7 mm thick at midpoint. Sepals 2, 8 mm long, ca 6 mm wide, 3.5 mm deep, ovate and somewhat concave, acute at apex, subcoriaceous, densely pubescent adaxially. Petals connate, subcoriaceous; tube 4.5 mm long, adnate at base to toms, with appressed and erect hairs adaxially, appressed-pubescent abaxially; lobes 4, 12.5 mm long, 4.5 mm wide at base, linear, tapering to an acute apex, with both appressed and short erect hairs adaxially, appressed-pubescent abaxially. Stamens 1.5-1.8 mm long; connectives cushion-shaped, erect-pubescent. Carpels ca 1.3 mm long [immature]; ovaries obscured by their long appressed hairs; style apex hispidulous. Fruit unknown.

This species, known only from the fragmentary type collection, is similar to D. corona tum, but differs in its long-acuminate membranous leaves, longer pedicel, and hairy corolla. Additional collections are necessary to determine how consistently this suite of differences is maintained. The specimen, collected in March, was from a swamp forest in eastern Sumatra (Fig. 5).

7. Disepalum aciculare D. M. Johnson, sp. nov. (Figs. 1H, 2F, 6)

Habitu et florum aspectu Disepalo longipedi King simile, sed corollae lobis acicularibus, ad medium latioribus, ad apicem angustioribus, et stylis ad apicem crispato-pilosulis diversum.

Tree 3-12 m tall, with DBH of up to 50 cm; bark smooth, gray to dark gray. Leafy portion of twigs 0.8-3.4 mm thick, thinly sericeous at first, soon glabrate, yellowish, reddish, or olive-brown to gray or brown with epidermis exfoliating in patches, finely longitudinally striate or wrinkled. Lamina of larger leaves 11.3- 16.4 cm long, 4-5.4 cm wide, chartaceous to membranous, oblong to oblong- oblanceolate, base cuneate and decurrent on petiole, apex acuminate, the acumen 8-13 mm long, glabrous adaxially, sericeous abaxially when young, soon glabrate; midrib impressed adaxially, raised and keeled abaxially; secondary veins 9-14 per side, these and higher-order veins indistinct or slightly prominent on both leaf surfaces. Petiole 6-9 mm long, 1.2-1.5 mm wide, sparsely pubescent to

1989] JOHNSON; DISEPALUM 373

glabrate. Inflorescence terminal or leaf-opposed, 1- or 2-flowered; pedicels 3.9- 5.1 cm long, 0.6-1.2 m m thick at midpoint , rusty sericeous at first, then sparsely pubescent. Sepals 2, 6.5-9 m m long, 5-6 m m wide, chartaceous, broadly ovate, acute at apex, with t iny dense erect hairs adaxially, sparsely appressed-pubescent abaxially. Petals connate, subcoriaceous; tube 2-5 m m long, adnate to torus, sparsely appressed-pubescent on abaxial surface; lobes 4, 4-5 m m long, 0.5-0.6 m m wide at widest point, narrowed near middle with port ion distal to this point lanceolate and often with a longitudinal furrow, glabrate adaxially, sparsely appressed-pubescent abaxially, Stamens 1.2-1.4 m m long; connectives depressed- globose, densely erect-puberulent. Carpels ca 100-150; ovaries 0.6-0.9 m m long, densely sericeous; styles 0.5-0.6 mm long, with crisped hairs at apex; ovules 2. Torus 5 m m in diam. Pedicel in fruiting stage 4.5-5 cm long, 1.7-2.3 m m thick at midpoint , glabrate. Torus in fruiting stage 1.5-2.2 cm in diam, calyx and corolla persistent; carpophores 3.5-6.5 cm long, 0.7-0.9 m m thick at midpoint , obliquely clavate and 2.0-2.7 m m thick at apex. Monocarps 3 6 -1 1 0 + , concolorous with carpophores, 8 .5-13 m m long, 6.5-7.5 m m wide, ellipsoid, apex rounded or apiculate, base attenuate, glabrate; monocarp wall 0.2 m m thick. Seeds 2, rarely only 1, 7-7.5 m m long, 6-6.7 mm wide, irregularly ellipsoid to globose.

TYPE. BORNEO. SARAWAK: [probably Mt. Matang], [late 1865 or early 1866], Beccari P. B. 756 (HOLOTYPE: K!; ISOTYPES: G!, P!).

Phenology and distribution: Apparent ly confined to low elevation forests in the vicinity o f Kuching, Sarawak, East Malaysia (Borneo) (Fig. 5), D. aciculare has been collected in flower in March, May, September, and October, and in fruit in September and October.

Color notes: F rom the few data available, the flowers and fruits seem similar in color to those o f other sympetalous disepalums: the sepals are green, the anther connectives are red, and the carpophores are pink.

Additional specimens examined: BORNEO. SARAWAK: Simanggang Dist., Sg. Eutalang, Anderson 13118 (K, L, SING); Kuching, 1893, Bartlett s.n. (BM); Simanggang, Division 2, Brooke 10753 (L); 31/2 mi from Kuching, various dates, Haviland = 1652 (BM, CAL, GH, K, P, SING); Kuching, Oct 1905, Hewitt s.n. (BM); Simunjun Distr., Simunjan F. R., Murthy 11732 (SING); without definite locality, Native Collector 477 (A); Stapok Forest Reserve, Kuching, Nitta & Yoshida 1 (KYO); Se- mengoh Forest Reserve, Kuching, Sinclair 10193 (A, K, SING, US).

D i s e p a l u m aciculare is most similar to D i s e p a l u m longipes; both are medium- sized trees with flowers with moderate to large numbers o f carpels; the corolla lobes o f D. aciculare, however, are needle-like, linear-lanceolate, widest near the middle and tapering to the apex, with scattered appressed, pale hairs, while those of D. longipes are spatulate or clavate, widest near the apex, with more or less spreading rusty hairs. The hairs at the apices o f the styles are also different: crisped in D. aciculare, rigid and straight in D. longipes. The two species are allopatric.

8. DISEPALUM LONGIPES King (Figs. 1 I, 2G, 6D, H)

Disepalum longipes King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61: 69. [13 Jun] 1892. TYPE: MALAYStA: Johore, Gunong Panti, Jun 1880, King's Collector 231 (HOLOTYPE: CAL, photograph!; ISOTYP~: B!, BM!, G!, K!).

Tree 1.5-8 m tall with a DBH of 5-10 cm. Leafy port ion o f twigs 0.8-4.6 m m thick, sericeous at first, soon glabrescent, reddish or yellowish-brown to grayish- brown, longitudinally wrinkled or finely striate, occasionally with decurrent ridges f rom the petiole base. Lamina o f larger leaves 10-22.5 cm long, 3.3-6.5 cm wide, chartaceous, oblong-oblanceolate, cuneate and decurrent at base, apex acuminate, the acumen 6-16 m m long, glabrous adaxially, initially sericeous abaxially, soon sparsely appressed-pubescent; midrib impressed adaxially, raised abaxially; sec-

374 BRaTTONIA [VOL. 41

b

l C

g i g, c , d . . . . . . . J

FIG. 6. Disepalum aciculare D. M. Johnson. A. Sepal, adaxial view. B. Flower. C. Close-up of corolla lobe, adaxial view. D. Close-up of corolla lobe of Disepalum longipes, adaxial view, for comparison. E. Leaf, adaxial surface. F. Aggregate fruit. G. Stamen, adaxial view. H. Style of Disepalum longipes for comparison with I. I. Carpel, lateral view with portion of ovary wall cut away to show attachment of ovules. J. Monocarp. K. Seed, view of exostomal end. L. Seed, lateral view. A-C, E, and G-I are from Haviland 1652 (K), D is from Ridley 4183 (SING), F is from Sinclair & bin Tassim 10193 (SING), and J-L are from Native Collector 477 (A).

ondary veins 9-17 per side, these and higher-order veins indistinct to slightly raised on both surfaces. Petiole 4 -10 m m long, 0 .7-1.6 m m wide, sparsely appressed-pubescent . Inflorescence terminal, somet imes leaf-opposed, l - or 2-flowered; peduncles 0.8-2 m m long, pedicels 2.4-6.2 (7.2) cm long, 0.6-1.3 m m thick at midpoint , sparsely appressed-pubescent . Sepals 2, 6.5-10.5 m m long, 5.5-8 m m wide, subcoriaceous, ovate to orbicular, with dense short erect hairs adaxially, sparsely appressed-pubescent abaxially. Petals connate, subcoriaceous; tube 2 .5- 5 m m long, broadly obconic, adnate to torus, sparsely appressed-pubescent abaxially; lobes 4, 3.5-6.5 m m long, 0.7-1.5 m m wide at widest point, narrowly spatulate, blunt at apex, sparsely appressed-pubescent except on adaxial apical pad, which has dense minu te erect hairs. S tamens 1-1.5 m m long; connect ives depressed-globose, densely erect-puberulent . Carpels 36--ca 60; ovaries 0.4-1 m m long, sparsely to densely sericeous; style 0.5-0.9 m m long, setulose at apex; ovules 2. Torus 3.5-6 m m in d iam. Pedicel in fruiting stage 3.8-4.7 cm long, 0.8-1.5 m m thick at midpoint , sparsely pubescent. Torus in fruiting stage 7-13 m m in


diam, sepals and petals persistent; carpophores 2.2-4 cm long, 0.8-1.5 m m thick at midpoint , t runcate and 1.5-2.5 m m thick at apex. Monocarps 18-47, concolorous with carpophores , 9 -13 m m long, 7 .5-10 m m wide, ellipsoid, rounded at base and apex, glabrate; m onoca rp wall 0.2 m m thick. Seeds 2, rarely 1, 8-8.5 m m long, 7.5 m m wide, ellipsoid.

Phenology and distr ibution (Fig. 5): D i s e p a l u m longipes has been collected in flower and fruit throughout the year on the Malay Peninsula; f rom Sumat ra it is known by flowering collections f rom June and August only. It occurs at elevations of 300 to 500 mete rs in the Malay Peninsula, where it appears to be c o m m o n only on the sandstone ridges and s u m m i t o f Gunung Panti in southern Johore. The species occurs at elevations o f 850 to 1000 meters in Sumatra .

Color notes: Flower and fruit colors for D. longipes m a y be summar ized as follows: calyx and corolla green, anther connectives red, rose-red, or deep red, gynoecium cream-colored; fruits light p ink to red, borne on red carpophores.

Additional specimens examined: MALAY PENINSULA. DINDINGS: Bruas, road to Sungei Rotan, Curtis 3437 (K, SING). JOHORE: Gunong Panti Forest Reserve, Ahmad S.328 (A, C, G, K, KLU, L, SING); forest around the Norwegian Field Station ca 15 km j'4W Kota Tinggi, Bremer & Bremer 1826 (KLU); Kota Tinggi, Gunung Panti, on the Panti Ridge somewhere between E and W Peak, Chin 2165 (A, KLU, L, MO); Summit Gunong Panti Barat, Cockburn FRI 7847 (KEP); Gunong Panti, Corner SFN 26170 (K, KEP, NY, SING); summit ridge between G. Panti E and G. Panti W, Panti F. R., Everett FR1 13809 (A, L); G. Besar massif, below G. Pukin above S. Segamat, Lubis F. R., Everett FRI 14013 (A, KEP); Gunung Panti, Holttum SFN 18071 (SING); Gunung Panti E, Koehum- men 29052 (KEP); Kota Tinggi Dist., Panti F. R., Matan F, D. 53931 (KEP); summit of Gunong Panti, Maxwell 81-135 (SING); Kluang Dist., Kluang F. R., Ng KEP 98082, 98083 (KEP); Ridley 4183 (BM, CAL, SING, UC); Gunong Panti East, Shah & AhmadMS.2916 (SING); Gunong Panti, ShukorAS.3 (A, C, KLU, L, SING); ShukorAS.30 (C, SING); NW Gunong, Upper Camp, Whitmore FRI 8815 (KEP, L). SUMATRA. W. Kust. Loeboek Sikaping, Boschproefstation hr. C. C. 7658 (K); Afd. Again Brani, Biinnemeijer 3210 (K, U); Taram, E of Pajakumbuh, sandstone region of River Tjampo, Maradjo 304 (L).

D i s e p a l u m longipes has been regarded, virtually f rom its original description, as conspecific with D. a n o m a l u m , and the two are indeed s imilar in aspect o f the foliage and flowers. The two species differ consistently, however , in habit, details o f the corolla and gynoecium, and ecology. D i s e p a l u m longipes is more often a tree, reaching eight meters in height, while D. a n o m a l u m is a treelet reaching only three meters. The corolla lobes o fD. longipes are less broadly spatulate than those of D. a n o m a l u m , often tending toward clavate, and are sparsely covered with ferruginous hairs; the lobes in D. a n o m a l u m are often strongly rhombic at the apex, with a dense covering o f ferruginous hairs. Finally, D i s e p a l u m longipes is exclusively montane , while D. a n o m a l u m occurs only in lowland white-sand areas.

9. DISEPALUM ANOMALUM Hook. f. (Figs. 1J, 2H)

Disepalum anomalum Hook. f., Trans. Linn. Soc. London 23: 156, pl. 20. 1860. TYPE: BORNEO: Lowlands of SARAWAK, in 1857, Lobb s.n. (HOLOTYPE, K!).

Disepalum grandiflorum Ridley, Bull. Misc. Inform. 1912: 384. 1912. TYPE: BORNEO. SARAWAK: Baram, Hose 142 (LECTOTYPE, designated here: K!; ISOLECq'OTYPES: B!, BM!).

Small tree 1.5-3 m tall. Leafy por t ion o f twigs 0.7--4 m m thick, reddish-brown to pale yellowish-gray, soon becoming light gray with epidermis exfoliating, slightly longitudinally wrinkled, glabrous. L e a f l amina 7.5-15.2 cm long, 2.4-5.6 cm wide, chartaceous to subcoriaceous (rarely membranous-char taceous) , elliptic to oblong, occasionally oblanceolate to obovate , base cuneate and short decurrent, apex acuminate with a blunt acumen 4-13 m m long, glabrous adaxially, at first sericeous abaxially, soon sparsely pubescent; midr ib impressed adaxially, raised and keeled abaxially; secondary veins (5) 8-12 per side, these and higher-order veins slightly raised on both surfaces. Petiole 4-1 1 m m long, 0 .8-1.7 m m wide,

376 BgaTTO~aA [VOL. 41

glabrous. Inflorescence leaf-opposed or sometimes terminal, I- or 2 (3)-flowered; pedicels (2.7) 3.7-6.5 cm long, (0.4) 0.7-1.2 m m thick at midpoint , glabrous to sparsely pubescent. Sepals 2, 5-12 m m long, (3.5) 6.5-1 1 m m wide, chartaceous, orbicular or broadly ovate, rounded or acute at apex, with fine erect pubescence adaxially, glabrous or glabrescent abaxially. Petals connate, subcoriaceous; tube 1.5-11 m m long, broadly obconic to narrow and abruptly flaring, adnate to torus, densely pubescent abaxially; lobes 4, 3-8 m m long, 0 .6-2 m m wide at narrowest point, 1.6-2.3 m m wide at widest point, spatulate, apex blunt, hairy adaxially except on spatulate or rhombic apical portion, uniformly pubescent abaxially. Stamens 1.5-2.1 m m long; connectives cushion-shaped, densely erect-puberulent. Carpels ca 30; ovaries 0.8-1 m m long, densely sericeous; styles 0.9-1.4 m m long, short-pubescent to hispidulous at apex; ovules 2, rarely 1. Torus 2.5-6.5 m m in diam. Pedicel in fruiting stage 1.9-5.2 cm long, 0.7-1.6 m m thick at midpoint , glabrous. Torus in fruiting stage 5.5-12 m m in diam, calyx and corolla persistent; carpophores 0.6-2.5 cm long, 0.7-1.2 m m thick at midpoint , truncate and 1.1- 1.5 m m thick at apex. Monocarps 7-26 (53), concolorous with carpophores, 9- 12 m m long, 7-9 m m wide, ellipsoid, apex rounded, sometimes apiculate, base rounded, glabrate; monocarp wall 0.1 m m thick. Seeds 2, rarely 1, 7-8.5 m m long, 6-6.5 mm wide, ellipsoid.

Phenology and distribution: Disepalum a n o m a l u m is known from Brunei and Sarawak on Borneo, almost entirely from coastal kerangas forests at elevations o f 30 to 400 meters (Fig. 5). Flowering material o f this species has been collected in all months except March and September; fruiting collections are known from February, August, November , and December.

Color notes: The flowers have green sepals, green to yellowish-white petals, and buf f to yellowish-white anther connectives. The color o f the connectives probably reflects immature flowers. The monocarps are pink to purplish black.

Additional specimens examined: BORNEO. BRLrNEI: Badas F. R., Ashton BR UN 5558 (K, L); Anduki F. R., Corner BRUN 5350 (L, SING); Teraja Forest Reserve, Seria District, enroute from Kpg. Mendaram to Bt. Teraja (W route), Hotta 12795 (KYO, L); Bt. Teraja, Teraja Forest Reserve, Seria Dist., Hotta 12865 (KYO); en route from Bukit Teraja to Kpg. Mendaram (E route), Teraja Forest Reserve, Seria District, Hotta 12933 (KYO); Belait District, Badas, van Nie14102 (MO, L; both sheets mixed with D. coronatum); Andalau Forest Reserve (west), Sinclair & Kadim bin Tassim 10455 (A, B, L, SING, US). SARAWAK: Baram District, Marudi, Hose 214 (K; lectoparatype ofD. grandiflorum); Marudi, 17 Jun 1895, Hose[?] s.n. (BM); without definite locality, Native Collector 1836 (US); Marudi (Claudetown), IV Division, Richards 1007 (K); Matang, Ridley 12255 (K, SING); Kuching Dist., 6th Mile F. R., Yakup 7723 (K, L, SING).

Disepalum a n o m a l u m appears to be sympatric with D. coronatum; both occur in the white-sand kerangas forests o f Sarawak and Brunei. Van Niel 4102 f rom Brunei is a mixed collection of the two species, suggesting that they were found at the same site. The two species are readily separated in flower or fruit by the corolla lobes: hairy and spatulate in D. anomalum, nearly glabrous and linear in D. coronatum. There is also a tendency for the twigs o f D. a n o m a l u m to be pale gray, while those o f D. coronatum are dark gray or gray-brown.

Disepalum a n o m a l u m may also co-occur with D. aciculare, but again morphology o f the corolla lobes will separate the two. In addition, D. a n o m a l u m is a shrub or small tree, and has only 30-55 carpels per flower and carpophores 0 .6- 2.5 cm long; D. aciculare is a medium-sized tree with 100-150 carpels per flower and carpophores 3.5-6.5 cm long. There is a tendency for the leaves in D. anomalum to be, as Hooker described them, " 'pellucid-punctate," while those o f D. aciculare are smooth, but this is not a constant difference. Differences between D. anornalum and D. longipes are discussed under the latter.

The types o f D. a n o m a l u m and D. grandif lorum represent two extremes o f


flower size n o w we l l - connec ted by m o r e recent col lect ions. Th i s p h e n o m e n o n of var iab le flower size was also obse rved in D. coronatum and , to a lesser extent , in D. platypetalum; i n D. anomalum, however , it is no t a c c o m p a n i e d by va r i ab i l i ty in n u m b e r of corol la lobes.

A c k n o w l e d g m e n t s

I t h a n k the cura to rs of the fol lowing he rba r i a for m a k i n g spec imens ava i lab le to me for this s tudy: A, B, BM, BO, C, CAL, F, G, G H , H B G , IBSC, K, KEP, K L U , K Y O , L, M I C H , MO, MSC, N-Y, P, PE, S I N G , U C , a n d US. I t h a n k B. Verdcour t for select ing specific spec imens at K for me to bor row. T r a n s l a t i o n s of labels wr i t ten in Ch inese were p r o v i d e d for m e by C.-H. Tsou , B. C. Tan , a n d T. K o y a m a . B. M e u r e r - G r i m e s t rans la ted cri t ical passages in G e r m a n for me. R. C. Ba rneby corrected the La t in diagnosis . W. R. A n d e r s o n a r ranged the t ransfer o f the bu lk of m y loans f rom M I C H to N Y in order tha t I migh t comple t e this s tudy. I t hank P. KeBler, a n d P. Maas a n d the A n n o n a c e a e t e a m at U t r e c h t for their careful a n d useful reviews of the manusc r i p t . N. A. M u r r a y suggested this project and also m a d e m a n y helpful suggest ions on the ma nusc r i p t .

L i t e r a t u r e C i t e d

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analysis of Tetrameranthus. Taxon 37: 346-353. Kubitzki, K. 1987. Origin and significance of trimerous flowers. Taxon 36:21-28. Le Thomas, A. 1980. Ultrastructural characters of the pollen grains of African Annonaccae and their

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revision of disepalum (annonaceae)

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