SYSTEMATIC STUDIES OF BORNEAN ZINGIBERACEAE V. ZINGIBEROIDEAE OF LAMBIR HILLS, SARAWAK

This paper reports the subfamily Zingiberoideae (Zingiberaceae) of Lambir Hills National Park, Sarawak, Malaysia. Twelve species representing Boesenbergia, Globba, and Zingiber are recorded. Systematic and ecological notes are provided, and the key to the Bornean species of Boesenbergia is updated. Four species, B. flabellata, B. ischnosiphon, B. lambirensis, and B. lysichitoides, and one variety, Z. longipedunculatum var. lambirense are described as new to science. The lectotype of B. grandifolia is designated. In addition, new combinations of Scaphochlamys and Haplochorema are reported.


INTRODUCTION
inflorescence is only found in the genus Boesenbergia, and all species from Lambir Hills have inflorescences of this type. For both types, flowers open from apex to base in the inflorescence. This mode of flowering is elsewhere in the family only known in the closely related Haplochorema and the probably more distantly related Caulokaem pferia Larsen (Smith, 1987).
An important character to identify species, first commented by , is anther dehiscence patterns. Thecae of some species open by terminal pores, but in others thecae dehisce throughout their length. Smith (1987) described that these pores are not "true pores" in most species, because usually slits are present on the lateral surface of each theca nearest to the style. However, at least in some species there is no slit on the inner face, as observed in B. flabellata (described below). All Lambir species have anthers dehiscing by pores, except B. ischnosiphon with thecae dehiscing about half of their length.
Descriptions of mature fruits and seeds of Bornean Boesenbergia are limited; only found in Poulsen (1993;B. urceoligena) and Cowley (1998;B. jangarunii). We describe mature fruits and seeds of B. ischnosiphon and B. lambirensis below. Seeds of the four species have a deeply laciniate aril as described for the Malay Peninsular species by Holttum (1950). They may be dispersed by ants as found in Bornean Globba (Pfeiffer et al., 2004). Holttum (1950) also mentioned the thin wall of the capsule, but the wall of B. ischnosiphon is not very thin, and does not agree with the other two Bornean species, B. lambirensis and B. jangarunii, or peninsular species, while the character is not described for B. urceoligena.
Notes -According to Smith (1987) no recent collection of the species had been seen. The type material contains no flowers on the type sheet and the original description by Ridley (1909: 57) is incomplete. Although the description says the lamina is glabrous, the type material has sparse long hairs on the upper surface and somewhat denser on the lower surface. The material from Lambir is similar to the type specimen in many characters such as 2-or 3-leaved shoots, obovate leaves with attenuate base of similar size, cuspidate and pubescent bracts, while ligules are shorter in Lambir plants (up to 2 cm,Fig. 2c) than in the type (2-4 cm). On the other hand, floral sizes of the Lambir plants are somewhat different from the description in having longer and glabrous corolla lobes (10-12 mm) rather than hairy ones of 6 mm (1/4 inch) described by Ridley (1909: 57). In Lambir plants anthers open by pores, but there is no description about the character in the original description. We tentatively identify the material from Lambir as B. hirta since no other materials of the species from the type locality are available at the moment.
Habitat & Ecology -Boesenbergia ischnosiphon often forms a dense clump with dozens of single-leaved shoots, on floors of wet and relatively open forests. Flowers are pollinated by halictid bees (Sakai et al., 1999, cited as B. gracilipes).
Etymology -The epithet 'ischnosiphon' refers to the plant's long and thin corolla tube.
Notes -Boesenbergia ischnosiphon is characterized by single-leaved shoots with a long petiole and an elliptic lamina. There are quite a few specimens from Borneo with these characteristics, but they are not identical. Burtt 8281 (E) from Mulu is different from Lambir material in having pubescent anthers dehiscing throughout their length, as well as some pubescence in the upper part of the bracts, sheaths and ligules. P.C. Boyce 272 and 297 (K) from Brunei have much shorter corolla tubes (c. 5.5 cm) than the material from Lambir (c. 9 cm). More collections from Borneo, including pickled flowers, are needed to examine the variation within species.
The collection Burtt 8281, mentioned above, was initially misidentified as Boesenbergia gracilipes (K. Schum.) R.M. Sm. by Smith (1987). Whereas Burtt 8281 is still in concordance with the description of Boesenbergia, a closer inspection of the inflorescence of the type of B. gracilipes indicates that this should be transferred to Scaphochlamys (see p. 110).
Habitat & Ecology -Most fertile shoots of B. lambirensis bear only one leaf; occasionally none. The first author found a similar plant, but with a several-leaved shoot (S. Sakai 1013, KYO), at the third waterfall of Lambir Hills where B. lambirensis and B. hirta grow in close proximity. Since young leaves of the plant had some purple on the lower surface similar to B. hirta, the plant might be a hybrid.
The species often form a loose clump at relatively open and wet sites. It flowers sporadically without particular flowering season (Sakai, 2000), and is pollinated by halictid bees (Sakai et al., 1999). In both publications, the species is cited as B. grandifolia. The fruit of B. lambirensis has a thin wall. Seeds occasionally germinate while the fruit is still attached to the infructescence but only after it has dehisced.
Note -In Smith (1987) Burtt & Woods 2430 was identified as B. grandifolia (ut B. grandiflora). But the material was quite different from the lectotype (see below) in having much smaller and elliptic leaves (26 cm long) rather than huge, obovate leaves (80 cm long, including petiole in B. grandifolia) with attenuate bases. In addition, it  Perennial herb c. 50 cm tall (Fig. 5e), fertile shoot 2-4-leaved, rarely single, usually covered with a few bladeless sheaths at the base, especially in young shoots; bladeless sheath to 17 by 3 cm, papery, but usually much shorter, purplish even after it has dried, densely hairy outside, glabrous inside; lamina 40-48 cm long, c. 13 cm wide, obovate, plain green, purplish towards base and petiole, basal part of midrib on the upper surface dark brown, below with long appressed hairs especially on and around midrib, almost glabrous on the upper, hairs getting denser toward base and apex, apex acute or slightly acuminate, base attenuate into thick petiole with groove or winged; petiole to 3 cm long, densely covered with robust hairs; ligule fragile and short-lasting, to 2 (?) cm long, probably 2-lobed to the base, densely pubescent outside, almost glabrous inside; sheath densely pubescent, margin membranous. Inflorescence terminal, with c. 20 flowers arranged in a one-sided spiral (Fig. 6e); sterile bract a few, c. 7.5 by 1.3 cm (outermost), coriaceous, narrowly ovate, apex of the outermost sterile bracts exceeding the fertile bracts by c. 1.5 cm, densely pubescent outside, glabrous inside, apex obtuse and acuminate; fertile bracts 2-3.5 cm long, to 4 mm wide, papery to coriaceous, oblong, densely pubescent with long hairs outside, glabrous inside; bracteole c. 3 by 1 cm, lanceolate, softly pubescent outside, glabrous inside, margin membranous, apex acute (Fig. 6b). Flower borne singly, white (Fig. 6a); calyx c. 10 mm, unilaterally fissured for half its length, densely pubescent outside, glabrous inside, apex 2-toothed (Fig. 6c); corolla tube c. 3.8 cm, pubescent outside, glabrous inside except pubescence at the throat; corolla lobes membranous, lanceolate, glabrous, dorsal lobe c. 15 by 6 mm, laterals c. 12 by 4 mm; lateral staminodes c. 9 by 5 mm, oblong to obovate, glabrous; labellum c. 16 by 16 mm, broadly ovate, slightly 3-lobed at the apex, white with a central yellow line with red blotches at the throat, glabrous (Fig. 5f, 6a); stamen 9 mm long, shortly pubescent; filament c. 2 mm long, shortly and sparsely pubescent; anther c. 3.5 mm long, open by pores from which a slit along the internal side of each theca, glabrous; anther crest c. 4 mm, lateral part of the crest swollen to enfold the stigma, glabrous; stigma cup-shaped with apical opening, with sparse hairs around the mouth (Fig. 6d); ovary c. 4.5 mm, probably incompletely trilocular, a few ovules per locule, axial placentation, pubescent only upper half; epigynous gland c. 5 mm, in two parts, linear, apex blunt. Capsules unknown. Distribution -Borneo, Sarawak, Lambir Hills.
Habitat & Ecology -The plant is rather abundant on marshy ground of Sg. Liku, Lambir Hills, but has not been collected from other sites.
Etymology -The epithet refers to the vegetative characters and habitat of the species resembling Lysichiton (Araceae). Gastrochilus grandifolius Valeton (1914) 241; 98. -Boesenbergia grandifolia (Valeton) Merr. (1921)  Notes -The original description (Valeton, 1914: 241) did not cite specimens precisely, but mentioned "Borneo, collected by Nieuwenhuis?". Later,  cited Nieuwenhuis 936 and 939 as the species, which are kept at the Herbarium Bogoriense. The two are from the same locality, the former has a 2-leaved sterile shoot, and the latter a single-leaved shoot with an inflorescence. We selected the latter as the lectotype (Fig. 7b).

Boesenbergia parva (Ridl.) Merr.
Boesenbergia grandifolia is more or less glabrous in the vegetative part. The species is further characterized by large leaves measuring 70-100 cm including the petiole.

KEY TO BORNEAN BOESENBERGIA
(modified from Smith, 1987) 1a. Creeping herbs; shoots normally single-leaved; inflorescence more or less sessile; anther dehiscing by slits . The genus Haplochorema was erected by K. Schumann based on the unilocular ovary with basally affixed ovules. However, this character is not diagnostic for the genus, because the ovary may also be trilocular or incompletely partitioned Smith, 1987), and it is now well-understood that the number of locules varies within Boesenbergia and other genera (Burtt & Smith, 1964). In the current system the only character to distinguish Haplochorema from Boesenbergia is the labellum being deeply bilobed or emarginate, flatly held rather than saccate. Smith (1987)  Note -  placed the species in Gastrochilus, because the ovary of the species is trilocular, which is not a diagnostic character of the genus any more. According to the label information on the type and original description by  the flower is "Kaempferia-like" with a deeply bilobed labellum and a broad, recurved anther crest, but it is difficult to confirm this examining the poorly preserved flowers on the type.

SCAPHOCHLAMYS Baker
The genus is distinguished from allied Bornean genera (Boesenbergia, Kaempferia, and Haplochorema) (Smith, 1987) by having flowers in cincinni and short, free, basal spurs on the thecae. Notes -This species, originally described under Haplochorema, was moved to Boesenbergia by Smith (1982). She saw a specimen from Mulu (Burtt 8281, E) with reproductive characters of Boesenbergia, and she thought the specimen and the type material of H. gracilipes at the Herbarium of the Royal Botanic Gardens Kew were identical, failing to notice a small inflorescence under the tag. A well-developed and clearly visible lax inflorescence with flowers in cincinni, as noted in the description by Schumann (1904), and spiral arrangement of bracts found in the material at the Herbarium of Museo di Storia Naturale dell'Università, Firenze (FI) (Fig. 7a), further indicates that the species belongs to Scaphochlamys, which was misunderstood and not accepted by Schumann (1899Schumann ( , 1904 (Smith, 1987). Most specimens identified to be B. gracilipes by Smith (1988) probably belong to Boesenbergia.
The lax inflorescence of S. gracilipes indicates affinity with S. polyphylla among Bornean Scaphochlamys, but the former is different from the latter in the more delicate inflorescences and single-leaved fertile shoots. After Smith (1988) revised the Bornean Zingiber including 16 species, 14 species have been described by Theilade & Mood (1997a, b, 1999 and Theilade & Christensen (1998). It is difficult to update the key provided by Smith (1988) without embarking on a proper revision of the genus. Perennial herb 1-1.5 m tall, up to 21 leaves per shoot. Lamina up to 27 by 6.5 cm, oblong to lanceolate, apex acuminate to 1.5 cm, base cuneate, glabrous above, sparsely hairy below; petiole short to 5 mm long, pubescent; ligule to 3 mm long, shallowly 2-lobed, more or less hairy, margin thinner, sometimes ciliate; sheath sparsely hairy to glabrous but with somewhat dense hairs below ligules near the margin, margin otherwise membranous. Inflorescence radical, c. 5.5-10 by 2.5-4.5 cm, ovoid, with a peduncle of up to 40 cm, prostrate on the ground, peduncular scales to 6 cm long, tubular at the base up to 1.2 cm, pubescent on both surfaces, hairs are more sparse on inner surface; bracts up to 4 by 1.8 cm at the base of inflorescence, oblong, hairy on both surfaces, denser at the base of the outer surface, lower margin membranous, apical margin densely ciliate especially when young, apex obtuse and occasionally very shortly acuminate and ciliate, not reflexed when alive, although rarely slightly open outwards when dried (Fig.  7d); bracteole c. 26 by 7 mm, oblong, open to the base, pubescent outside, glabrous inside except ciliate apex. Flower borne singly, white; calyx c. 22 mm long, tubular, unilaterally fissured in the upper half, pubescent outside in the lower 5 mm, otherwise glabrous, apex acute or 2-lobed for 3 mm, minutely ciliate; corolla tube 35-40 mm long, glabrous outside, hairy at the throat inside; corolla lobes glabrous, laterals c. 26 by 4.5 mm, linear, connate to the labellum and each other for basal c. 8 mm, dorsal c. 25 by 10 mm, lanceolate; labellum c. 20 by 17.5 mm, 3-lobed, central lobe c. 15 by 10 mm, ovate, shallowly 2-lobed at the apex, side lobes 4 by 3 mm, oblong and entire, glabrous; anther with c. 1 mm long filament, thecae c. 12 mm long, 2.5 mm wide each, glabrous, dehiscing throughout their length, anther crest c. 8 mm long, triangular, enfolding the style; style c. 6 cm long, glabrous, stigma c. 0.7 mm wide, cup-shaped fringed by long hairs, placed slightly higher than the tip of the anther crest; ovary c. 4 mm, hairy; epigynous glands c. 7 mm, linear, 2-lobed. Capsules unknown. Distribution -Borneo, Sarawak, Lambir Hills.
Habitat & Ecology -The species is often found abundant in swampy secondary forests. Bright red inflorescences are very conspicuous on the forest floor. The white flowers open around noon and are frequently visited by medium-sized Amegilla bees collecting pollen (Sakai et al., 1999).
Note -Zingiber longipedunculatum is a species characterized by the long, prostrate peduncle and strongly reflexed bracts, which can easily be recognized in herbarium sheets, although size and shape of inflorescences and bracts vary significantly. In the type and most herbarium collections examined from Borneo, the inner surface of the bract is glabrous. Plants from Lambir Hills have rather flat, not reflexed, bracts, pubescent on both surfaces, and deviate from other materials.
Notes -We have not seen the types of Z. odoriferum or the variety.  described the arachnoid indumentum on the lower surface of the leaves, although it could be variable, and the fusiform spike of c. 20 cm supported by a 40-100 cm long peduncle. These characters more or less agree with the Lambir material.
The species was described by Blume on the basis of collections from Java, and Valeton recognized four varieties, three from Java and one from Borneo. Valeton noted that the Bornean variety differed in the more narrow, acute inflorescence.