The Marantaceae of Sabah, Northern Borneo

The Marantaceae of Sabah, Northern BorneoAuthor(s): Karen Clausager and Finn BorchseniusReviewed work(s):Source: Kew Bulletin, Vol. 58, No. 3 (2003), pp. 647-678Published by: Springer on behalf of Royal Botanic Gardens, KewStable URL: http://www.jstor.org/stable/4111147 .Accessed: 18/10/2012 03:15

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp

.JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range ofcontent in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new formsof scholarship. For more information about JSTOR, please contact [email protected].

.

Royal Botanic Gardens, Kew and Springer are collaborating with JSTOR to digitize, preserve and extendaccess to Kew Bulletin.

http://www.jstor.org

http://www.jstor.org/action/showPublisher?publisherCode=springer

http://www.jstor.org/action/showPublisher?publisherCode=kew

http://www.jstor.org/stable/4111147?origin=JSTOR-pdf

http://www.jstor.org/page/info/about/policies/terms.jsp

KEW BULLETIN 58: 647 - 678 (2003)

The Marantaceae of Sabah, northern Borneo

KAREN CLAUSAGER' & FINN BORCHSENIUS'

Summary. A taxonomic treatment is provided for the Marantaceae of the Malaysian state of Sabah in northern Borneo. Thirteen species are recognised, four of which are described as new to science: Phacelophrynium aurantium, Phacelophrynium laxum, Phrynium grandibracteatum and Phrynium stenophyllum. One new combination is made: Stachyphrynium placentarium (Lour.) Clausager & Borchs. Keys to the genera and species are provided together with illustrations of the four new species.

INTRODUCTION

The Marantaceae is a family of 31 genera and c. 550 species of small to moderate- sized herbs distributed throughout the tropics, with the exception of Australia. About half the genera and c. 450 species are Neotropical while the remaining genera and c. 100 species occur in the Palaeotropics. Only a few members of the

family extend into the temperate regions of South and North America (Andersson 1998). The family is one of the most derived families within the order Zingiberales (Kress 1990). Several species are of economic importance, such as Maranta arundinacea as a high-quality starch crop and Calathea spp. for ornamental plants.

While several studies have been carried out on the taxonomy of neotropical Marantaceae (e.g., Andersson 1977, 1981, 1986; Kennedy 1978, 1986) only sporadic accounts exist for SE Asia, in spite of the fact that members of the Marantaceae often form an important constituent of the herb layer in SE Asian rain forests. Moreover, existing taxonomic treatments are mostly quite old and lack keys and contemporary descriptions of the species. Important references include the family monograph by Schumann (1902), as well as local treatments of the Marantaceae in the Malay Peninsula (Ridley 1899, Holttum 1951), Borneo (Ridley 1906) and the Philippine Islands (Ridley 1909). No substantial contribution to our knowledge of the

taxonomy of the SE Asian Marantaceae has been made in the last 50 years. Here we present a taxonomic revision of the Marantaceae in the Malaysian state of

Sabah in northern Borneo. When the study was initiated, only seven species were known from Sabah; we have identified 13, four of which are described here as new. We believe that this is symptomatic of the state of knowledge of the Marantaceae in Borneo and the Malesian region in general. Given the relatively wide distribution of most of these species, the keys and descriptions should be useful to taxonomists in most of the western Malesian region.

Accepted for publication April 2003. 1 Department of Systematic Botany, Institute of Biological Sciences, University of Aarhus, Herbarium, Build. 137

Universitetsparken, DK-8000 Aarhus C, Denmark.

647

648 KEW BULLETIN VOL. 58(3)

The study is based on existing material in the herbaria A, AAU, BM, C, K, KEP, L, P, SAN, SING, UKM and Sabah Parks (abbreviations according to Holmgren et al. 1990) as well as our own collections. Generic concepts follow Andersson (1998); nomenclatural rules used in connection with the relevant sections of the descriptions are those of the "Saint Louis Code" (Greuter et al. 2000). A complete list of the material examined is available on request.

INFLORESCENCE STRUCTURE AND DESCRIPTIVE TERMINOLOGY

According to Andersson (1976, 1981, 1998) the inflorescence in Marantaceae is a polytelic synflorescence in which the basic unit (florescence) is an open, spiciform or capitate thyrse. This is made up of an indeterminate (or rarely determinate) main axis carrying a number of distichously or spirally arranged bracts, each subtending a highly specialised partial florescence (florescence component), derived from a branched system of determinate axes (repeatedly enriched, two- flowered cymules). Due to condensation of the cymule axes the flowers are borne in pairs in which one flower forms a mirror image of the other. Solitary flowers or flower triplets are known in a few genera, but none of these occur in Sabah. The florescence component may include from 1 - 20 flower-pairs, though the most common number found is 2 - 4.

A radically different interpretation of the Marantaceae inflorescence has, however, been presented by Kunze (1983, 1985, 1989) and Kirchoff (1986). According to these authors the flower-pair itself constitutes the florescence, with each flower representing a single-flowered cyme derived from the partial florescence type commonly found in other Zingiberales, including Canna, the sister group of the Marantaceae (Kress 1990, Andersson & Chase 2001).

In view of the continuing controversy concerning the correct interpretation of the Marantaceae inflorescence we have here preferred to use a neutral descriptive terminology, avoiding terms such as florescence, florescence component, cymule etc., the typologically correct application of which remains uncertain. We thus describe the inflorescence as consisting of a main axis plus a number of branches of

increasing order. If there are no branches (as in Stachyphrynium latifolium) we describe the inflorescence as simple. Bracts are referred to by indication of their position in the inflorescence and the structure they subtend. For the specialised flower-bearing branch systems (florescence components sensu Andersson) we use Kunze's term "special paraclade", as this would correctly signal their nature irrespective of which typological interpretation is favoured. Special paraclades can be dolichoblastic, i.e., with elongate axes (Donax, Schumannianthus) or brachyblastic with strongly condensed axes (Phacelophrynium, Stachyphrynium, most Phrynium species). By the term synflorescence we refer to the entire flower-bearing part of the inflorescence, i.e., all parts excluding the peduncle. A schematic presentation of the

terminology used is given in Fig. 1.

THE MARANTACEAE OF SABAH, NORTHERN BORNEO 649

B

Bract Prophyll

Fertile bract Flower

Special paraclade Fertile bract

Interphyll

Fl(;. 1. A schematic drawing of the synflorescence of a Phacelophrynium-species. The shown example has four lateral branches with 3 - 5 special paraclades each subtended by distichous bracts, and the main axis has four special paraclades subtended by spirally arranged bracts. B schematic drawing of a special paraclade with three flower pairs and the associated bract, prophylls and interphyll. The determinate axes are truncated and thus appear indeterminate. In reality the axes are much more condensed and the synflorescence appear much more compact. In some Phrynium-species the bracts of both the main axis and lateral branches are spirally arranged.


DISTRIBUTION AND ECOLOGY

The five genera of Marantaceae represented in Sabah, Donax, Schumannianthus, Phacelophrynium, Phrynium and Stachyphrynium are all widely distributed in SE Asia. Donax, Schumannianthus and Phrynium occur throughout mainland Asia and the west-Malesian region, from India to New Guinea. Stachyphrynium is centred in mainland Asia with a centre of diversity in Thailand and Indochina, while Phacelophrynium is largely restricted to the Malesian region. Of the thirteen species found in Sabah, five appear to be endemic to Borneo (Phacelophrynium aurantium, P. laxum, Phrynium grandibracteatum, P stenophyllum and Stachyphrynium borneense). Stachyphrynium borneense, however, appears to be closely related to and perhaps conspecific with S. sumatranum, distributed in the more western parts of the Malesian region. The remaining eight species (Donax canniformis, Schumannianthus dichotomus, Phacelophrynium maximum, Phrynium hirtum, P pubinerve, P. villosulum, Stachyphrynium latifolium and S. placentarium) are all widespread in SE Asia.

Seven species are widespread in Sabah. Four species (Phrynium hirtum, P stenophyllum, Stachyphrynium placentarium and Phacelophrynium laxum) appear to be confined to the western part, and one (Phrynium grandibracteatum) has so far only been recorded from eastern Sabah. Only Phacelophrynium laxum grows in mountain forest above 1000 m elevation. We have not recorded any species as associated with ultramafic soils or limestone outcrops, as is known in several other families. Some species, such as Stachyphrynium borneense, are found in complete shade on the floor of primary forest, whereas others, such as Phacelophrynium aurantium and P maximum, seem to prefer to grow next to streams; others, such as Donax canniformis and Phrynium pubinerve are capable of growing in more open and disturbed habitats. Most species can, however, be found both in primary and secondary forest, and none appears to be highly sensitive to moderate disturbance of the forest structure.

KEY TO THE GENERA OF MARANTACEAE IN SABAH

1. Stems aerial, richly branched ................................... 2 Stems subterranean (rosulate plants) ............................. 3

2. Flower less than 2 cm long; lamina up to 30 x 20 cm ............. 1. Donax Flower 2 - 4 cm long; lamina 10 - 15 x 5 - 10 cm ....... 2. Schumannianthus

3. Flower with 1 outer staminode ..................... 3. Phacelophrynium Flower with 2 outer staminodes .................................. 4

4. Sepals at least half as long as the corolla tube; corolla tube usually equalling the lobes or shorter ................................... 4. Phrynium

Sepals less than half as long as the corolla tube; corolla tube at least twice the length of the lobes ............................ 5. Stachyphrynium

TAXONOMY

1. Donax Lour., Fl. Cochinch.: 11 (1790); Holttum, Gard. Bull. Singapore 13: 266- 271 (1951); Rolfe,J. Bot. 45: 242 - 244 (1907); Andersson in Kubitzki, The Families and Genera of Vascular Plants IV: 289 (1998). Type: Donax arundastrum Lour.


Actoplanes K. Schum. in Engler, Das Pflanzenreich IV, 48: 33 (1902). Lectotype (here designated): Actoplanes canniformis (G. Forst.) K. Schum.

Tall herbs, up to 5 m or more, with richly branched aerial stems. Synflorescence richly branched, lax, the branches narrowly spiciform. Special paraclades markedly dolichoblastic, with 1 - 2 flower pairs; interphylls absent; bracteoles 2 per flower pair, small, glandular. Corolla tube equalling the lobes or shorter; staminal tube only slightly longer than corolla tube, inconspicuous; outer staminodes 2, petaloid; ovary 3-locular. Fruit indehiscent; seeds exarillate. Three to six species ranging from Thailand and Malaysia to the New Hebrides (Andersson 1998). One species in Sabah.

1.1. Donax canniformis (G. Forst.) K Schum., Bot.Jahrb. Syst. 15 (4): 440 (1893).

Thalia canniformis G. Forst., Fl. Ins. Austr.: 1 (1780). Type: New Hebrides, Forster s.n. (BM!).

Actoplanes canniformis (G. Forst.) K. Schum. in Engler, Das Pflanzenreich IV, 48: 34 - 35 (1902).

Donax arundastrum Lour., Fl. Cochinch.: 11 (1790). Type: Cochinchina, Loureiro s.n. (BM!).

Maranta grandis Miq., Fl. Ind. Bat. Suppl.: 616 (1860). Type: Indonesia, Sumatra, Teijsmann s.n. (BO?).

Donax grandis (Miq.) Ridl., J. Straits Branch Roy. Asiat. Soc. 32: 176 (1899).

Branched herb, up to 5 m tall or more. Leaf sheath 8 - 20 cm long; petiole absent; pulvinus 1 - 1.5 cm long, green, slightly pubescent; lamina 8 - 30 x 4 - 20 cm, upper surface dark green, lower surface pale green, with hairs along the prominent midrib. Inflorescences several on each plant, richly branched, lax, composed of up to 15 major branches of increasing orders, to 30 cm long; each branch subtended by a 3 - 6 cm long, brown, papery bract and bearing a bicarinate prophyll, 2 - 4 cm long, just above its base; distal internodes of the inflorescence branches elongate, 1.5 - 3 cm long; each node with a brown, papery bract, 23 - 32 mm long, subtending either a prophyllate branch or a special paraclade with 1 - 2 flower pairs; primary axis of special paraclades 25 - 30 mm long, pedicel of individual flowers 5 - 25 mm long, associated bracteole c. 2 mm long, fleshy. Flower white, 15 mm long; sepals subulate, 2.8 x 0.8 mm; petal lobes subulate, 8.3 x 2.8 mm; corolla tube 4.9 mm long; staminal tube 1.8 mm longer than corolla tube; outer staminodes slightly unequal, lobes 5.6 - 6.9 x 1.4 - 2.1 mm; cucullate staminode 4.2 x 2.8 mm; callose staminode with large petaloid lobe 5.6 x 3.5 mm, cavity 1.4 x 1.4 mm; stamen 4.4 mm long, with a small, narrow, petaloid appendage 2.4 mm long; style 4.2 mm long, curved, stigmatic cavity 0.8 mm diameter; ovary 1.4 mm long, pubescent. Fruit green to white, c. 1 cm in diameter, completely smooth and globose, indehiscent; seeds 1 - 3.

DISTRIBUTION. Peninsular Malaysia, Borneo, Sumatra, Java and the Philippines. Donax canniformis is found throughout Sabah at elevations up to 800 - 1000 m.

HABITAT AND ECOLOGY. Primary and secondary forest, often in relatively disturbed habitats along streams and rivers, or along railways and roads.


COMMON NAMES. Bamban (Malay); lias (Dusun). SELECTED SPECIMENS EXAMINED (from a total of 25). SABAH: Kampong Brungis,

Tuaran (Jesselton W. C.) Forest Distr. Cuadra 1262 (A); Beluran Distr., Sapapayau Forest Reserve, Amin et al. 111544 (SAN); Tenom, near Pangi, Kokawa & Hotta 2625 (L); Tenom Distr., Near Kampong Pomila'an, Gibot 66805 (SAN); Sepilok Forest Reserve, Sandakan, Kadis 2575 (A); Sandakan Distr., Bt. Tingkar, Joseph 56759 (SAN); Sandakan Suan Lambah, Cuadra 1462 (SAN); Bettotan, Castro 3195 (A); Keningau Distr., Ulu Biah, Sundaling 83918 (SAN); Marotai, Maidin 2677 (A); Sandakan, Myburgh Province, Elmer 20255 (A); Telupid Distr., Kg. Lalau Kapuk Keramuak, Mansus et al. 108856 (SAN); Kalabakan Distr., NBT logged area, km 27 Luasong road, Krispinus 95643 (SAN); Keningau Distr., Sook plain F. Res., Sundaling 80772 (SAN); Kalabakan Distr., Ulu Segama, Krispinus 95447 (SAN); Kalabakan Distr., Hap Seng logging area, km 19 Ulu Sg. Toa, Pikkoh 101443 (SAN); Imbak R., Poulsen & Kjeldsen 1611 (AAU).

NOTES. The material examined has been determined as either Donax canniformis or D. grandis (Miq.) Ridl., but it all belongs to the same species. Holttum (1951) speculated that the two names might be synonymous; in fact Merrill had already synonomised the two in 1924 (Merrill 1924). The nomenclatural history of the genus Donax is confusing; see Schumann (1893, 1902), Ridley (1899), Rolfe (1907) and Holttum (1951).

2. Schumannianthus Gagnep., Bull. Soc. Bot. France 51: 169 - 177 (1904); Holttum, Gard. Bull. Singapore 13: 271 - 273 (1951); Rolfe, J. Bot. 45: 244 (1907); Andersson in Kubitzki, The Families and Genera of Vascular Plants IV: 289 (1998). Type: Schumannianthus dichotomus (Roxb.) Gagnep.

Tall, branched herbs. Corolla tube half the length of the lobes or less; outer staminodes 2, petaloid. Fruit dehiscent; seeds arillate. Two species, one in Sri Lanka and India (Malabar), another in Malaysia, Indochina and the Philippines (Andersson 1998). One species in Sabah.

2.1. Schumannianthus dichotomus (Roxb.) Gagnep., Bull. Soc. Bot. France 51: 176 (1904); Holttum, Gard. Bull. Singapore 13: 272 - 273 (1951).

Phrynium dichotomum Roxb., Asiat. Res. 11: 324 (1810).

Tall herb with branched aerial stem. Leaf sheath 5 - 6 cm long, green, glabrous; pulvinus 5 - 10 mm long, green, glabrous; lamina 7 -9 x 3- 5 cm, green, glabrous. Inflorescence terminal, lax, spiciform, 8 - 10 cm long, with 2 - 4 bracts each subtending at least one flower pair. Flower up to 4 cm long, white.

DISTRIBUTION. Malaysia, Indochina and the Philippines. HABITAT AND ECOLOGY. Mostly in disturbed areas, including roadsides. COMMON NAME. Bemban (Brunei). SPECIMEN EXAMINED. SABAH: Tenggilan, Tandom 2833 (L). NOTES. Schumannianthus dichotomus looks very similar to Donax canniformis, but

can be distinguished by its smaller leaves, larger flowers, less branched


inflorescence, and dehiscent fruit. The generic distinction has, however, been questioned by Andersson (1998), and the whole Donax/Schumannianthus complex is much in need of revision. We have seen only a single collection of S. dichotomus from Sabah, (annotated Donax canniformis) and for this reason our description is incomplete. The nomenclature of Schumannianthus is closely linked with that of Donax; for a thorough account we refer to Rolfe (1907) and Holttum (1951).

3. Phacelophrynium K. Schum. in Engler, Das Pflanzenreich IV, 48: 120 (1902); Holttum, Gard. Bull. Singapore 13: 294 - 296 (1951); Andersson in Kubitzki, The Families and Genera of Vascular Plants IV: 288 (1998). Lectotype (here designated): Phacelophrynium maximum (Blume) K Schum. [Basionym: Phrynium maximum Blume].

Rosulate ground herbs. Inflorescence a branched synflorescence; bracts persistent, usually distichously arranged on the inflorescence branches. Special paraclades markedly brachyblastic, with 2 - 4 flower pairs; interphylls usually present; bracteoles absent. Sepals of the same length as the corolla tube or longer; corolla tube equalling lobes or shorter; outer staminode 1, petaloid; ovary 3-locular. Fruit dehiscent; seeds arillate. Six to nine species distributed from Thailand and the Malay Peninsula eastwards to Borneo and the Philippines; a single species has been described from New Guinea (Andersson 1998). Three species in Sabah.

NOTES. No type was designated by Schumann when he described the genus (Schumann 1902; Greuter et al. 1993). We have chosen Phacelophrynium maximum (Blume) K. Schum. as the lectotype, as this agrees well with Schumann's original protologue for the genus.

KEY TO THE SPECIES OF PHACELOPHRYNIUM

1. Upper leaf surface with distinct lighter markings along the midrib ? ?

3.2. P. laxum Upper leaf surface uniformly green ............................... 2

2. Synflorescence bracts bright orange; sheath, petiole and lower bracts with dark hairs; plant less than 1 m tall ...................... 3.1. P. aurantium

Synflorescence bracts brown to green; sheath, petiole and lower bracts glabrous; plant up to 3 m tall ............................. 3.3. P. maximum

3.1. Phacelophrynium aurantium Clausager & Borchs. sp. nov. Phacelophrynio maximo similis, sed statura minori ad 1 m non 3 m alta, bracteis clare aurantiis non viridibus vel brunneis, vaginis, petiolis, pedunculis et bracteis infimis pubescentibus non glabris differt. Typus: Malaysia, Sabah, Danum Valley Field Centre, Orchid Trail, 5 Dec. 1999, Clausager 19 (holotypus AAU!; isotypi K!, KEP!, SAN!, Universiti Malaysia Sabah!).

Rhizomatous ground herb, up to 1 m tall, usually forming small clusters. Leaves 4 - 7 per shoot, distichous; sheath 28- 39 cm long, brown to green, covered by stiff, brown to black hairs; petiole 3 - 31 cm long, light green, with scattered hairs similar to those on the sheath; pulvinus 2 - 4 cm long, light green, glabrous; lamina oblong-


elliptic, 20 - 37 x 8 - 12 cm, dark green above, pale green below, glabrous. Inflorescence interfoliar, erect; peduncle 10 - 19 cm long, green, with scattered brown hairs; synflorescence congested, 8 - 10 x 5 - 6 cm; first order branches 3 - 5, erect, to 6 cm long, arranged in two opposing rows forming an angle of c. 120', the proximal branch usually with 1 - 2 shorter, second order branches at its base; fertile bracts 10 - 20 per branch, arranged in a pattern similar to that of the branches themselves, but distichy often imperfect due to the twisting of the plane of insertion between subsequent pairs of bracts; terminal 4 - 5 cm of the main inflorescence axis with 8 - 15, spirally arranged, fertile bracts; all bracts ? similar, 2 - 4 cm long, lanceolate to oblong, orange, with a ? dense cover of blackish hairs; flower pairs 3 - 4 per special paraclade, associated prophylls c. 25 x 8 mm, interphylls c. 5 x 2 mm. Flowers white, 25 - 30 mm long; sepals subulate, 10 x 1 mm; petal lobes subulate, 11 x 3 mm, translucent, slightly orange at the tip; corolla tube 10 - 14 mm long; staminal tube 4 - 10 mm longer than the corolla tube; outer staminode with a petaloid lobe 3 x 2.3 mm; cucullate staminode 4.5 x 3 mm, dotted with small brown spots; callose staminode 6 x 6 mm, cavity 2.7 x 2 mm; stamen c. 3 mm long, with a narrow petaloid appendage, 2.5 x 0.7 mm; style with free part c. 4 mm long, curved, stigmatic cavity 1.3 mm diameter; ovary 2 mm long. Fruit globose, green to brown, c. 10 mm in diameter; seeds 1 - 3, oblong, 7 - 8 mm long, brown, aril white, bifid, 6 mm long. Fig. 2.

DISTRIBUTION. Known from Sabah east of the Crocker Range massif, and from eastern Sarawak. Apparently endemic to the island of Borneo.

HABITAT AND ECOLOGY. Found in primary forests in relatively shaded locations, usually near streams or rivers. It grows in lax clusters with 10 or more shoots. The flowers are visited and pollinated by large trap-lining bees of the genus Amegilla. An individual flower lives for just one day, but the synflorescences are able to produce flowers continuously for at least 3 months with an average of 4 flowers open per day (pers. obs.). These observations agree well with reports from Sarawak on the pollination ecology of Phacelophrynium maximum, Stachyphrynium cylindricum and S. griffithii (Kato 1996, Momose et al. 1998).

COMMON NAME. Kobu pelandok (Dusun). SPECIMENS EXAMINED (other than the type). SABAH: Tenom Valley, hills behind

Agricultural Station, Clausager 11 (AAU, SAN); Lahad Datu distr., Ulu Sg. Danum, near Kuala Sg. Segama, Stone et al. SAN85189 (L); Danum Valley, km 63, Maikin & Lidah 131127 (SAN); Danum Valley, W trail, Blewett 3 (UKM); Danum Valley Ulu Segama, along trail VJR Forest, Lantoh SAN109675 (KEP, SAN); Danum Valley, Segama W trail, Madani SAN112946 (KEP, SAN); Danum Valley Field Centre, Palum Tambun R., Lee 124807 (SAN); Danum Valley, W trail, Andrews 729 (UKM); Lahad Datu distr., Ulu Sungei Segamat, Stevens et al. 499 (KEP); Sepulut distr., Labang, Sepulut Forest Reserve, Fidilis SAN125289 (KEP); Keningau distr., Sepulut, Sungai Saburan,J impin SAN106981 (K); Telupid distr., N Margin Tawai Forest Reserve, Meijer SAN131944 (A); Tawau Hills National Park, Dolois et al. 09001 (Sabah Parks); Tawau Hills National Park, trail to Sulphur Spring, Clausager 22 (AAU, SAN); Tawau Hills National Park, Johannsen 12 (AAU, SAN); Tawau, St Lucia, F. D., Kadir A2054 (A); Sibuga-Labuk road mile 5, Sandakan, Meijer 05611 (Sabah Parks); Tabin Wildlife


cm

cuc os If

B? A

sta

E . .....

... ..... .. ... c a cal

o -]mm

i

?

J

r •oo_•10 cm

-. 4

FIG. 2. Phacelophrynium aurantium. A habit; B lower part of shoot showing leaf sheaths, petiole, peduncle and

synflorescence; C lamina of leaf; D flower; E fruit with remnants of calyx. Drawn from holotype byJens Christian Schou. Labels for flower parts: s sepal; p petal; os outer staminode; cal callose staminode; cuc cucullate staminode; sta fertile stamen.


Reserve, 4.2 km from HQ, Borchsenius 531 (AAU, SAN); Secondary forest near Imbak R., Poulsen 1602 (AAU). SARAWAK: 7th division, Ulu Sg. Apa, between Bkt. Goram and Bkt. Bakak, Chai 36205 (KEP); 4th division, Dulit Range, downstream Sg. Sirui, Dyg. Awa & Yii P C. 46718 (K); Ulu Sg. Semawat, Belaga SAR43721 (SAN); Upper Rejang R., Clemens & Clemens 22137 (K); Sungei Mayeng, Tau Range, Purseglove 5302 (K).

NOTES. Phacelophrynium aurantium differs from all other species in the genus in its bright orange bracts and hairy sheaths, petioles and lower bracts. Almost all collections of this species have previously been identified as P maximum (Blume) K. Schum., which, however, has glabrous sheaths, petioles and bracts, and is much larger than

P. aurantium. Furthermore, it has pale green to brown bracts, never

orange ones as in P aurantium.

3.2. Phacelophrynium laxum Clausager & Borchs. sp. nov. ab omnibus aliis speciebus Phacelophrynii lamina secus costam flavovirenti-striata, synflorescentia in duabus vel tribus partibus divisa; bracteis viridibus; floribus luteis distinguenda. Typus: Malaysia, Sarawak, 7th division, Ulu Belaga, Sungai Semawat, C. Hansen 619 (holotypus C!; isotypi KEP!, SAN!).

Rhizomatous herb, up to 80 cm tall. Leaves 1 - 3 per shoot, distichous; sheath 8 - 13 cm long, glabrous, green; petiole 13 - 33 cm long, glabrous, green; pulvinus 3 - 4 cm long, glabrous, light green; lamina 15 - 30 x 8 - 15 cm, ovate, glabrous, the upper surface green with a pale yellowish green stripe along the midrib, lower surface pale greyish green with dark green veins, weakly plicate along the primary veins. Inflorescence interfoliar, erect; peduncle 14 - 20 cm long, green; synflorescence elongate, lax, 6 - 9 cm long; first order branches 1 - 2, simple, distichous, each subtended by a narrow, acuminate, green bract, 3 - 6 cm long; basal branch widely separated from the rest of the synflorescence by a 4 - 5 cm long internode; inflorescence branches 1 - 2 cm long, with 2 - 4, distichous fertile bracts, 1.5 - 2 cm long, light green; main axis terminated in a 1 - 2 cm long portion carrying 2 - 6 spirally arranged, fertile bracts, similar to those on the branches; flower pairs 2 - 3 per special paraclade, associated prophylls 10 - 15 mm long, interphylls c. 10 mm long. Flowers yellow, c. 1.5 cm long; sepals 10.5 x 1.8 mm; petal lobes 6.8 x 3 mm; corolla tube c. 6 mm long; staminal tube c. 3 mm longer than corolla tube; outer staminode with petaloid lobe 6.8 x 6 mm; cucullate staminode 3.8 x 1.5 mm, with small triangular appendage (trigger); callose staminode with a large petaloid appendage, 4.5 x 4.1 mm; stamen 2.3 mm long; style with free part 3.3 mm long, curved, stigmatic cavity 0.7 mm in diameter; ovary 2.3 mm long, densely covered in silky hairs. Fruit greenish brown, globose, c. 0.5 in cm diameter; seeds 1 - 3, brown, globose, 4 mm long; aril white, bifid, 1.5 mm long. Fig. 3.

DISTRIBUTION. So far known only from western Sabah and Sarawak. HABITAT AND ECOLOGY. Found in shade in mountain forests. The type specimen

was collected in hill dipterocarp forest. In Sabah the species has been collected on relatively steep slopes in disturbed forest in the Crocker Range area.

SPECIMENS EXAMINED (other than the type). SABAH: Road Kota Kinabalu - Tambunan km 39, Clausager & Borchsenius 1 (AAU, SAN); Road Kota Kinabalu -


cuc PP

] mm

cal

10 cm.

I p

s

A/

,\\

.. .. .....-] l m

yi

FIG. 3. Phacelophrynium laxum. A habit; B synflorescence; C flower. Drawn from holotype by Jens Christian Schou. Labels for flower parts: s sepal; p petal; os outer staminode; cal callose staminode; cuc cucullate staminode; sta fertile stamen.


Tambunan km 39, Johannsen & Poulsen 16 (AAU, SAN); N Borneo, G. Macara Fagal, Amdjah 145 (L). SARAWAK: 7th division, Ulu Belaga, Sungai Semawat, Hansen 920 (C); Gunung Mulu National Park, camp east of Melinau Gorge, Hansen 241 (C).

NOTES. Phacelophrynium laxum differs from other species of the genus in its lax synflorescence that consists of two or three parts separated by relatively long internodes. This, together with the yellow flowers and the lighter stripe along the midrib on the adaxial side of the leaf, makes it easily distinguishable from all other

Phacelophrynium species. In 1945, Bakhuizen van den Brink annotated a Phacelophrynium specimen as R

laxum Bakh. f. msc., but this name was never published. The specimen, collected in 1912 by Amdjah in North Borneo (now Sabah), shows the same synflorescence structure as the species described here. As holotype we have, however, preferred to select a specimen collected by Carlo Hansen in 1981 in Sarawak, as this includes well-preserved, pickled flower material. The epithet laxum refers to the structure of the synflorescence with its long internodes.

3.3. Phacelophrynium maximum (Blume) K. Schum. in Engler, Das Pflanzenreich IV, 48: 122 (1902).

Phrynium maximum Blume, Enum. P1. Javae 1: 37 (1827). Type: Indonesia, Java, Blume s.n. (holotype L!).

Phrynium tapirorum Ridl., Trans. Linn. Soc. London, Bot. III: 382 (1893); J. Asiat. Soc. Bengal: 180 (1899). Lectotype (selected by Holttum 1951): Malaya, Tahan R., Pahang, Ridley 2398 (K!).

Phacelophrynium tapirorum (Ridl.) K. Schum. in Engler, Das Pflanzenreich IV, 48: 122 (1902).

Phacelophrynium bracteosum Warb. ex K. Schum. in Engler, Das Pflanzenreich IV, 48: 123 (1902); synon. nov. Holotype: Philippines, Mindanao, Warburg 14615 (Bt); lectotype, (here designated): Philippines, Luzon, Albay province, Vidal 3976 (K!).

Rhizomatous ground herb, up to 3 m tall, usually forming dense clusters. Leaves 2 - 5 per shoot; sheath 38 - 80 cm long, green to brown, with sparse indumentum of dark hairs; petiole 26 - 180 cm long, green, glabrous; pulvinus 6 - 10 cm long, glabrous, yellowish green; lamina elliptic, 38 - 100 x 17 - 66 cm, glabrous, green above, pale green below, the rachis yellowish green. Inflorescence interfoliar, erect, sometimes appearing to protrude from the sheath of an accompanying leaf; peduncle 29 - 70 cm long, glabrous, green; synflorescence congested, 12 - 20 x 5 - 15 cm; first order branches 3 - 6, erect, to 11 cm long, distichously arranged, the

proximal branches usually with a few, shorter, second order branches; fertile bracts

up to 20 per branch, arranged in 2 rows forming an angle of 120 - 140 degrees, ovate, 3 - 4 x 2 - 2.5 cm, yellowish green to pale brown, reddish or dark brown; main inflorescence axis terminating in a 6 - 10 cm long portion carrying 10 - 15

spirally arranged fertile bracts, similar to those on the lateral branches; flower pairs 3- 4 per special paraclade, associated prophylls 2.5 x 1 cm, interphylls c. 2 x 0.6 cm. Flowers white or orange, c. 30 mm long; sepals subulate, 9 x 1.8 mm; petal lobes 14 x 4 mm; corolla tube c. 12 mm long; staminal tube c. 10 mm longer than corolla


tube; outer staminode with a petaloid lobe 7.5 x 3 mm; cucullate staminode 2.3 x 1.5 mm; callose staminode 9 x 6.8 mm; stamen c. 3 mm long; style with free part 4.7 mm long, curved, stigmatic cavity c. 1 mm in diameter. Fruit c. 10 mm long, globose to ellipsoid, sparsely hairy, green to brown, dehiscent; seeds 3, 7 - 8 mm long, brown, with a white translucent, bifid aril 5 mm long. Fig. 4.

DISTRIBUTION. Widespread in SE Asia, from the Malay Peninsula to Sumatra, Java, and Borneo.

HABITAT AND ECOLOGY. Seven of the specimens examined were noted specifically as growing near streams or rivers, and it appears that P maximum is more prevalent in marshy areas or near running water.

COMMON NAMES. Kebo-kebo, daun kukop, kebuntakang (Dusun). SELECTED SPECIMENS EXAMINED (from a total of 39). SABAH: Rd. to Danum

Valley Field centre, 2.5 km from field station, Clausager & Borchsenius 4 (AAU, SAN); Rd. Kota Kinabalu-Tambunan km 39, Clausager & Poulsen 8 (AAU, SAN); Lahad Datu Distr., FR. Ulu Segama, Lideh & Donggop 116974 (KEP, SAN); Tenom Distr. Ulu Tomani, 3 mile from Kg. Kaang, Madani 90839 (L); Ranau Distr., Lohan, Amin &

Jarius 114279 (KEP); Ranau Distr., rd. from Lohan to Mamut Copper mine near Tank 80, Beaman 10606 (A); Ranau Distr., Biton Tampias, Amin & Donggop 93943 (L); Kinabalu National Park, Boundary Rentis, Stevens et al. 649 (L); Kinabalu National Park, Dallas, Clemens & Clemens 27204 (K); Kinabalu National Park, Sayap substation, Poulsen & Borchsenius 1576 (AAU, SAN); Tongod Distr., Sg. Pantagaluang, Keramuak, Sundaling 96992 (L); Pembangunan Sabariah, SDN. Berhd. Sg. Keratam, Gibot 97066 (L); Lamag Distr., Ulu Sg. Lokan, Aban & Petrus 90661 (K); Sungai Meliau track, Telupid, Sandakan, Zainudin et al. 4822 (SAN, UKM); Trail from Bangbangan Camp to Maliau Falls, Maliau Basin, Poulsen &

Banggilon 1650 (AAU, SAN); 1 km up Kuala Burong, Tabin Wildlife Reserve, Poulsen & Abdilla 1683 (AAU, SAN); Tawau Hills Park, Johannsen 9 (AAU, SAN).

.7411

All,? 7 -vI I C[L

FIG. 4. Phacelophrynium maximum. A habit; B synflorescence. Photos F. Borchsenius.


NOTES. Phacelophrynium maximum is the largest species in the genus, reaching up to 3 m in height. Of the Marantaceae in Sabah, only Donax canniformis and Phrynium pubinerve grow equally tall. Its synflorescences are likewise among the largest of any in the genus. Leaves of P maximum are used by the indigenous people of Borneo to wrap food or as roofing material (Christensen 1997).

Phacelophrynium maximum is somewhat variable both in size and in the colour of its bracts and flowers. There appear to be at least two morphological forms. The first, known from both the Crocker Range area in western Sabah and from the Tabin Wildife Reserve in eastern Sabah, has pale brown to beige bracts and orange flowers. It grows to a height of c. 1.5 m. The second form, known from Kinabalu Park, Sayap substation and Danum Valley, has green to brown bracts and white flowers and may reach 3 m in height. Distinction between the two forms is, however, not clear, and plants with intermediate characters have been encountered, e.g., A. D. Poulsen & R. Abdilla 1683, AAU! with both white and orange flowers in the same inflorescence. For the time being we therefore prefer to maintain the variation displayed within a single taxon of the species level.

Based on examination of the types and descriptions of P bracteosum and R maximum we have synonymised those two names in this work. Schumann (1902) separated the species on the basis of whether there was a leaf accompanying the inflorescence (P maximum), or only a bladeless sheath (P bracteosum). Field observations, however, show that inflorescences both with and without an

accompanying leaf are commonly found within the same individual.

4. Phrynium Willd., Sp. P1. I. 177 (1797) (nom. cons.); K. Schumann in Engler, Das Pflanzenreich IV, 48: 52 (1902); Andersson in Kubitzki, The Families and Genera of Vascular Plants IV: 287 (1998). Type: Phrynium capitatum Willd., nom. illeg. (Pontederia ovata L., non Phrynium ovata Nees & Mart. (1823); P rheedei Suresh & Nicolson) [=R pubinerve Blume].

Phyllodes Lour., Fl. Cochinch.: 16 (1790). Type: Phyllodes placentaria Lour.

Rosulate ground herbs. Inflorescence a branched synflorescence, often

congested and capitate; bracts usually spirally arranged; special paraclades markedly brachyblastic to slightly dolichoblastic, with 2 - 4 flower pairs; interphylls usually present; bracteoles absent. Sepals half as long as the corolla tube but usually much

longer; corolla tube c. half as long as the lobes, rarely longer; outer staminodes 2, petaloid; cucullate staminode with an appendage near apex; ovary 3-locular. Fruit dehiscent; seeds arillate. Around 20 species from India and Sri Lanka eastwards to New Guinea (Andersson 1998). Five species in Sabah.

KEY TO THE SPECIES OF PHRYNIUM

1. Synflorescence borne at the base of the plant on a very short peduncle ..... 2 Synflorescence terminal on a long peduncle, sometimes appearing to protrude

from the petiole of the accompanyingleaf ........................ 3 2. Leaves uniformlygreen above ..................... 4.4. P.stenophyllum


Leaves distinctly variegated above with dark green stripes on a light green background .................................. 4.5. P. villosulum

3. Synflorescence elongate, branches widely separated - - 4.1. P. grandibracteatum Synflorescence capitate, branches congested ........................ 4

4. Lower bracts nearly glabrous; fruits bright red; leaves glaucous or green beneath

.......*.................................... 4.3. P. pubinerve Lower bracts densely hairy; fruits orange to pale red; leaves red beneath .....

..*........... ............................... 4.2. P. hirtum

4.1. Phrynium grandibracteatum Clausager & Borchs. sp. nov. ab omnibus aliis speciebus generi Phrynii bractea ramum basalem synflorescentiae subtendenti longa angustaque, bracteis fertilibus biseriatim abaxialiter in axe insertis differt. Typus: Malaysia, Sabah, Tawau Hills National Park, trail to Sulphur spring and waterfall, 10 Dec. 1999, K. Clausager 20 (holotypus AAU!; isotypi KEP!, SAN!, Sabah Parks!).

Rhizomatous ground herb, up to 1.5 m tall. Leaves 3 - 6 per shoot; sheath 28 - 43 cm long, glabrous, red at base, papery, brownish; petiole 24 - 41 cm long, glabrous, fresh green in colour; pulvinus 3 - 7 cm long, glabrous, fresh green; lamina oblong- elliptic, 34 - 60 x 15 - 25 cm, glabrous, fresh green above, light green below. Inflorescence interfoliar, erect; peduncle 26 - 30 cm long, glabrous, green; synflorescence markedly elongate, 13 - 19 cm long; first order branches 2 - 3, simple or the basal one with a single second order branch, branches distichously arranged but all turned to the same side of the inflorescence; basal first order branch 4 - 6 cm long, subtended by a very conspicuous, narrow, elongate bract, 15 - 22 cm long, red at base, brown towards the tip, partly enclosing the entire synflorescence; subsequent lateral branch inserted 4 - 7 cm above the first, subtended by a similar but smaller bract, to 12 cm long; additional lateral branches (if present) inserted at increasingly shorter distances and subtended by increasingly smaller bracts; fertile bracts 4 - 6 per inflorescence branch, distichous, ovate, 2 - 2.5 x 1.5 - 2 cm, light green to light brown; main inflorescence axis terminated in a 4 - 5 cm long portion carrying 5 - 6 distichous, fertile bracts, similar to those on the lateral branches; flower pairs 2 - 3 per special paraclade, associated prophylls 18 - 22 mm long. Flowers white, c. 2.5 cm long; sepals subulate, 15 x 3 mm; corolla tube 9 mm long, lobes subulate, 10 x 4.5 mm, with a pink tinge; staminodial tube 3 mm

longer than corolla tube; outer staminodes each with a petaloid lobe 5.4 x 3 mm; cucullate staminode 6 x 3 mm, yellow at the tip; callose staminode with free part 7.5 x 6 mm; fertile stamen 3.8 mm long, with a narrow petaloid appendage, 3 x 1.5 mm; style with free part 6 mm long, curved, stigmatic cavity 0.9 mm in diameter; ovary 1 - 2 mm long. Fruits pink to rose, globose to triangular in cross section, 2 cm in diameter, with prominent ridges and furrows, dehiscent; seeds 3, c. 14 mm long, pink to orange, ridged, triangular in cross section; aril small, 3 - 4 mm long, white, translucent. Fig. 5.

DISTRIBUTION. Known only from a few collections in SE Sabah and the adjacent northern part of East Kalimantan.

HABITAT AND ECOLOGY. In Sabah we have found this species in shade in primary forest. The collection from East Kalimantan was made in secondary forest.


sP SO

osB

cuc

I mm os

D cal D

cuc

sta os

P E cal/ p

iO 1 1 I / / • 7

\5cm lIo cm

10 cm

A C

FIG. 5. Phrynium grandibracteatum. A lower part of shoot showing leaf sheaths, peduncle and synflorescence; B leaf

showing petiole, pulvinus and lamina; C synflorescence with conspicuous long narrow bract; D flower; E flower, mirror image of D; F fruit; G fruit in cross section. Drawn from holotype byJens Christian Schou. Labels for flower

parts: s sepal; p petal; os outer staminode; cal callose staminode; cuc cucullate staminode; sta fertile stamen.


SPECIMENS EXAMINED (other than the type). SABAH: Tawau Hills National Park, Johannsen 11 (AAU, SAN). EAST KALIMANTAN: KPC area Bengalon, plot P-2, Ambri et al. 1580 (SAN); 1577 (K, SAN).

NOTES. Phrynium grandibracteatum is distinguished from all other species in the genus by its long, narrow bract subtending the basal branch of the synflorescence (Fig. 5C), and by having the fertile bracts inserted in two rows abaxially on the supporting axis. The presence of two outer staminodes in each flower combined with long sepals and a short corolla tube place the species unambiguously in the genus Phrynium, in spite of its unusual inflorescence structure.

4.2. Phrynium hirtum Ridl., J. Straits Branch Roy. Asiat. Soc. 32: 181 (1899); Holttum, Gard. Bull. Singapore 13: 290 - 291 (1951). Lectotype (selected by Turner 2000): Malaya, Johore, Gunong Panti, H. N. Ridley s.n., Dec. 1892 (SING!).

Phrynium inflatum Merr., J. Straits Branch Roy. Asiat. Soc. 85: 164 (1922); synon. nov. Type: Malaysia, Sabah, Batu Lima near Sandakan, M. Ramos 1488 (holotype K!).

Rhizomatous ground herb, to 1.8 m tall. Leaves 6 - 7 per shoot; sheath 49 - 84 cm long, green with brown hairs; petiole 21 - 58 cm long, green, glabrous; pulvinus 4 - 6 cm long, greenish brown, glabrous; lamina 30 - 50 x 11 - 14 cm, green above, green to red below, glabrous. Inflorescence interfoliar, erect, apparently protruding from the petiole of the accompanying leaf; peduncle 41 - 125 cm long; synflorescence capitate, densely congested, 3 - 6 cm in diameter, made up by 3- 4 branches of several orders, each 2 - 3 cm long; bracts subtending the proximal branches large, 6 x 3 cm, partly enclosing the entire synflorescence; fertile bracts spirally positioned, 2 - 3 x 1 - 1.5 cm, brown, becoming shredded and fibrous with age; flower pairs 2 - 3 per special paraclade. Flowers (description based on Holttum 1951): sepals c. 15 mm long; petal lobes 10 x 3 mm; corolla tube 9 mm long; outer staminodes unequal, petaloid lobes 14 x 3 mm and 10 x 2 mm, respectively; cucullate staminode 7 mm long; callose staminode 8 mm long; fertile stamen 7 mm long; ovary densely covered with silky hairs. Fruit globose, c. 2 cm in diameter, orange to light red; seeds 3, pink, c. 8 mm long; aril light beige, 2 mm long.

DISTRIBUTION. Sumatra, Borneo, Malaya. HABITAT. Found in shade in relatively dense forest. SELECTED SPECIMENS EXAMINED (from a total of 8): SABAH: Tenom Distr.,

Lumaku F. Res., L. Madani et al. 132698 (SAN); Kinabalu National Park, Sayap substation, alt. 1000 m, A. D. Poulsen & E Borchsenius 1575 (AAU, SAN); Poring Hot Spring, trail back from Canopy walkway, K. Clausager 18 (AAU, SAN); Rd. Kota Kinabalu-Tambunan km 39, K Clausager & F Borchsenius 2 (AAU, SAN).

NOTES. Phrynium hirtum can be distinguished from P pubinerve, which it otherwise resembles, by the red lower leaf surface, hairy bracts and pale red fruits. Holttum (1951) suggested that Phrynium hirtum and Phrynium inflatum might be the same. We have seen both types and there is no doubt that the two are conspecific.

4.3. Phrynium pubinerve Blume, Enum. P1. Javae: 38 (1827); Turner, Gard. Bull. Singapore 46: 129 (1994). Type: Indonesia, Java, Blume s.n. (holotype L!).


Pontederia ovata L., Sp. Pl.: 288 (1753). Type: element in Rheede's Hort. Malab., 11: 67, t. 34 (1692).

Phrynium capitatum Willd., Sp. P1. 1: 17 (1797), nom. illeg., [superfluous name based on the same type as Pontederia ovata L.]

Phrynium rheedei Suresh & Nicolson, Taxon 35: 355 (1986) [new name for Pontederia ovata L.].

Phrynium malaccense Ridl., J. Straits. Branch Roy. Asiat. Soc. 32: 180 (1899). Lectotype (selected by Turner 2000): Malaya, Perak, Gunong Tunggul Dindings, H. N. Ridley s.n., March 1896 (SING!).

Rhizomatous ground herb 0.7 - 3.5 m tall, sometimes forming dense clusters up to several metres in diameter. Leaves 2 - 3 per shoot; sheath 20 - 100 cm long, green, with a pale grey, lanose indumentum; petiole 6- 187 cm long, green, with indumentum similar to that of the sheath; pulvinus 3 - 14 cm long, yellowish green, slightly compressed laterally; lamina uniformly green, 23 - 82 x 9 - 30 cm, glabrous, slightly plicate along the primary veins. Inflorescence interfoliar, erect, usually appearing to protrude from the petiole of the accompanying leaf; peduncle 31 - 172 cm long, green; synflorescence capitate, densely congested, 4 - 8 cm in diameter, made up by 5 or more branches of up to several orders; bracts subtending the proximal branches enlarged, 3 - 4 cm long, dark brown, partially sheathing the entire synflorescence, their distal part quickly withering and dissolving into a decaying fibrous mesh; fertile bracts spirally arranged, 1.5 - 2 cm long, pale yellowish green, quickly withering and becoming brown at the tip; flower pairs 2 - 4 per special paraclade; associated prophylls 1.5 - 2 cm long. Flowers pinkish white, c. 1.8 cm long; sepals subulate, 10 x 1.5 mm; petal lobes subulate, 7.5 x 3.5 mm, pink, deflexed and curled; corolla tube 4.5 mm long; staminodal tube 4 mm longer than corolla tube; outer staminodes slightly unequal, petaloid lobes 3 - 4.5 x 1.8 - 3 mm; cucullate staminode 3 x 2 mm, dotted with small brown spots; callose staminode 3.7 x 1.8 mm; stamen 3 mm long; style with free part 3 mm long, curved, stigmatic cavity 0.9 mm in diameter; ovary 3 - 4.5 mm long. Fruits bright red, elongate to triangular, c. 15 x 10 mm, dehiscent; seeds 3, grey, c. 13 x 7 mm, with a basal, translucent, bifid aril. Fig. 6A.

DISTRIBUTION. Widely distributed in SE Asia, from India to Burma, Thailand, the Malay Peninsuala, Sumatra, Java, Borneo and the Phillipines. Common in forested areas throughout Sabah.

HABITAT AND ECOLOGY. Phrynium pubinerve seems to prefer a moderate level of disturbance, and is often found growing next to streams, roads and trails, or even in the understory of old rubber plantations. Both flowers and mature fruits are often found in the same inflorescence, indicating that flowering occurs over a considerable time.

COMMON NAME. Kobu (Dusun). SELECTED SPECIMENS EXAMINED (out of 33 total). SABAH: Beluran Distr., Hap

Seng Bridge, Labuk, Sundaling 91084 (SAN); Along Jalan Kabili, Sepilok Forest Reserve, Sandakan, Sam 49917 (SAN); Mile 32 Labuk Rd., Sandakan, Stevens et al. 399 (SAN); Sandakan distr., VJR 45 D, Lungmanis, Dewol et al. 113678 (SAN); Sandakan distr., Hs. Sg. Sapi, Lassan 116818 (SAN); Above Kampung Segindai,


A 9~E~i~llI ee %~

FIG. 6. A Phrynium pubinerve, synflorescence. Photo F. Borchsenius; B Phrynium villosulum, the very distinctive

variegated leaves. Photo A. D. Poulsen.

Ranau, Amin & Good 100225 (SAN); Kinabalu National Park, Eastern ridge, Poring, Kokawa & Hotta 4963 (L); Kinabalu National Park, Along Sg. Mamut, near Poring, Kokawa & Hotta 4843 (L); Kinabalu National Park, Sayap substation, Poulsen & Borchsenius 1574 (AAU, SAN); Pinangah distr., Ulu Sg. Melikop, Mansus & Dewol 109237 (SAN); Km 41 Imbak Rd., Luasong, Kalabakan, Fidilis & Sumbing 95701 (SAN); Kalabakan distr., Ulu Sg. Luasong Sabah Foundation area, NBT camp km 37, Luasong, Fidilis & Sumbing 68787 (SAN); Mnt. Trusmadi, Nooteboom 1365 (L); Keningau Distr., Crocker Range National Park, near headquarters, Poulsen &

Kandaung 1722 (AAU, SAN); N Borneo, unknown locality, Amdjah 430 (L); Lahad Datu distr., Jalan Teck Guan, Takun Ulu Bakapit, Saigol 93114 (SAN); Logging rd to Danum Valley Field Centre, 4.1 km from the field station, Clausager & Borchsenius 3 (AAU, SAN); Tabin Wildlife Reserve, trail 1 from Mud Volcano to road, Clausager & Borchsenius 5 (AAU, SAN).

NOTES. Most of the herbarium material examined was annotated Phrynium capitatum, but this is an illegitimate name. The name P capitatum was applied by Willdenow in 1797 to an element from Rheede's Hortus Malabaricus from 1692, that had earlier been named Pontederia ovata by Linnaeus (1753, p. 288). Willdenow

rightly recognised that a part of this element belonged to the genus Phrynium; his choice of a new name was, however, illegitimate as the taxon should rightly have been named Phrynium ovatum. The latter epithet became unavailable in 1823 when it was used for a different species of Phrynium (P ovatum Nees & Mart.). To rectify the situation Suresh & Nicolson (1986) published the new name Phrynium rheedei for the species. However, as shown by Turner (1994), the species had already been

correctly described as Phrynium pubinerve by Blume (1827). Leaves of Phrynium pubinerve are used for wrapping food and for roofing by indigenous tribes in Borneo (Christensen 1997).


4.4. Phrynium stenophyllum Clausager & Borchs. sp. nov. Phrynio hirto similis, sed synflorescentiis basalibus brevissime (< 5 cm, non 40 - 125 cm) pedunculatis, foliis angustioribus (39 - 56 x 8 - 12 cm non 30 - 50 x 11 - 14 cm) differt. Typus: Malaysia, Sabah, Tenom, Bontomon, 19 Nov. 1999, K. Clausager 14 (holotypus AAU!; isotypi KEP!, SAN!).

Rhizomatous herb, up to 1.6 m tall. Leaves 4 - 8 per shoot; sheath 48 - 60 cm long, green, densely covered by whitish, lanose hairs; petiole 11 - 53 cm long, green with some whitish hairs; pulvinus 3 - 6 cm long, green, glabrous; lamina 39 - 56 x 8 - 12 cm, green above, below sometimes slightly red, with a fine rim of hairs along the midrib. Inflorescence interfoliar, borne on a very short peduncle at the base of the plant; synflorescence capitate, c. 4 cm in diameter, made up of 3 - 4 or more branches, 3 - 4 cm long; fertile bracts spirally positioned, 2.5 x 1 cm, becoming shredded and split into fibres with age; flower pairs 2 - 3 per special paraclade, associated prophylls c. 2 x 0.7 cm, interphylls c. 1.5 x 0.5 cm. Flowers white, c. 3 cm long; sepals subulate, 22 x 3 mm; petal lobes subulate, 9 x 3 mm, deflexed and curled; corolla tube c. 20 mm long; staminal tube of same length as corolla tube; outer staminodes unequal, with petaloid lobes of 9 x 3.8 mm and 1.5 x 3 mm respectively; cucullate staminode 3.6 x 3 mm; callose staminode with a large petaloid appendage, 8.3 x 7.5 mm; stamen 2.3 mm long, with a small, narrow petaloid appendage, 1.5 mm long; style with free part 3.5 mm long, curved, stigmatic cavity 0.6 mm in diameter; ovary 4 mm long, densely covered by silky hairs. Fruit globose, c. 2 cm diameter, seeds 3, light red, c. 1.5 cm long, with white, bifid aril. Fig. 7.

DISTRIBUTION. In Sabah known only from the type locality near Tenom. It is also known from Ng. Sumpa in Sarawak, and has been seen in Batang Ai National Park, Sarawak (A. D. Poulsen, pers. comm.).

HABITAT AND ECOLOGY. The type plant was found on a steep slope in a logged, disturbed forest with vegetation dominated by bamboo.

SPECIMENS EXAMINED (other than the type). SARAWAK: 2nd division, Nanga Sumpa, Christensen & Poulsen 1998 (AAU).

NOTES. The species mostly resembles Phrynium hirtum, from which it differs in its longer and more narrow leaves (39 - 56 x 8 - 12 cm vs. 30 - 50 x 11 - 14 cm in P hirtum), and its basal inflorescence that is borne on a very short peduncle (< 5 cm long vs. 40 - 125 cm in P hirtum).

4.5. Phrynium villosulum Miq., Fl. Ind. Bat. Suppl.: 616 (1860); Kennedy, Kew Bull. 41: 725 - 731 (1986). Type: Indonesia, Sumatra, Teijsmann 4490 (holotype BO n.v.; isotype? U!).

Heliconia triumphans Linden ex Rodigas, Ill. Hort. 29: 59, t. 448 (1882). Type: Plate in Ill. Hort. 29, t. 448 (1882).

Phrynium basiflorum Ridl., J. Straits Branch Roy. Asiat. Soc. 32: 182 (1899). Type: Malaysia, Negeri Sembilan, Ridley 10000 (holotype K!).

Phrynium basiflorum var. nobile Ridl.,J. Fed. Malay States Mus. 4: 79 (1909). Lectotype (selected by Kennedy 1986): Malaysia, Perak, Ridley 14036 (hololectotype K!; isolectotype SING!).


]cm ~I cm

os

v 1 mm

sta mm cut

os

)CUC

/s

10

.OS

10 c

,A

SB

FIG. 7. Phrynium stenophyllum. A habit; B lower part of shoot showing leaf sheaths and synflorescence; C leaf

showing pulvinus and lamina; D flower; E seed. Drawn from holotype byJens Christian Schou. Labels for flower

parts: s sepal; p petal; os outer staminode; cal callose staminode; cuc cucullate staminode; sta fertile stamen. One petal is hidden on the back of the flower and is not visible on the drawing.


Rhizomatous ground herb, up to 2 m tall. Leaves 2 - 3 per shoot; sheath 38 - 57 cm long, green, with some whitish hairs; petiole 67 - 102 cm long, green, with few hairs similar to those on the sheath; pulvinus 8 - 11 cm long, green, glabrous, the junction between petiole and pulvinus marked by a narrow, light green band; lamina 32 - 45 x 15 - 22 cm, upper surface green with distinct, dark green stripes along the major veins, lower surface light green to red. Inflorescence interfoliar, borne on a very short peduncle at the base of the plant; synflorescence capitate, 5 - 6 cm in diameter, made up of 4 - 5 branches, each to 5 cm long; fertile bracts spirally arranged, 2 - 2.5 cm long, white to cream coloured with pink tips when young, turning brown and becoming shredded distally when old; flower pairs 2 - 3 per special paraclade, associated prophylls c. 2 x 0.7 cm, interphylls c. 1.5 x 0.5 cm. Flowers white, c. 3 cm long; sepals subulate, 20 x 3 mm; petal lobes subulate, 13 x 3.8 mm, deflexed and curled, slightly pink at tip; corolla tube 16 mm long; staminodal tube 3 mm longer than corolla tube; outer staminodes unequal, with petaloid lobes of 4.5 x 1 mm and 10 x 9 mm, respectively; cucullate staminode 7.5 x 4.5 mm, dotted with small brown spots; callose staminode 3 x 3 mm, basally fused with the larger outer staminode; stamen 4.5 mm long, with a narrow, threadlike appendage; style with free part 6 mm long, curved, stigmatic cavity 1 mm in diameter; ovary 4.5 mm long, covered by silky hairs. Fruit globose, c. 1.5 cm diameter; seeds 1 - 2, c. 10 mm long, aril 5 mm long, white, translucent. Fig. 6B.

DISTRIBUTION. The Malay Peninsula, Sumatra, and Borneo. In Sabah, Phrynium villosulum has been found only in a few quite widely separated locations: Tenom and Poring in western Sabah, and Tawau in SE Sabah.

HABITAT AND ECOLOGY. Usually in habitats with a moderate level of disturbance, such as steep slopes next to waterfalls.

SELECTED SPECIMENS EXAMINED (from a total of 12). SABAH: Tawau ("Tawao"), Elphinstone province, Elmer 20945 (L); Tawau Hills Park, Johannsen 13 (AAU, SAN); Tenom, N ridge of Mt Malutut, about 15 km N of Tenom, alt. 450 m, Kokawa & Hotta 2443 (SAN); Kalang Waterfall, Tenom Valley, Clausager 13 (AAU, KEP, SAN); Tenom Valley, hills behind Agricultural Station, Clausager 10 (AAU, KEP, SAN); Poring Hot Springs, trail back from Canopy walkway, Clausager 17 (AAU, KEP, SAN); Maliau Basin, Agathis Camp, Poulsen 1656 (AAU, KEP, SAN).

NOTES. Phrynium villosulum is easily recognized by its distinctly variegated leaves that recall those of some neotropical Calathea species. According to Holttum (1951), the flower of P villosulum has two narrow outer staminodes and a large callose staminode. The flowers examined by us all had one narrow and one broad outer staminode, while the callose staminode was quite small and basally adnate to the larger of the two outer staminodes. We believe that Holttum's description reflects a misinterpretation of the floral parts rather than a real difference in flower morphology between Bornean and west Malaysian specimens of P villosulum.

Kennedy (1986) cited the type of Phrynium villosulum as "Teijsmann 4490 (BO)" with no indication of type status. However, there is a specimen of Phrynium villosulum in U, with number 4490, but no collector. It seems most likely that the two specimens are from the same collection by Teijsmann, and that the one in U is an isotype.


5. Stachyphrynium K. Schum. in Engler, Das Pflanzenreich IV, 48: 45 (1902); Holttum, Gard. Bull. Singapore 13: 273 - 279 (1951); Andersson in Kubitzki, The Families and Genera of Vascular Plants IV: 288 (1998). Lectotype (here designated): Stachyphrynium latifolium (Blume) K. Schum. (basionym: Phrynium latifolium Blume).

Rosulate ground herbs. Inflorescence simple or a sparsely branched synflorescence; bracts usually distichous, persistent; special paraclades brachyblastic, with 1 - 3 flower pairs; interphylls present; bracteoles absent. Sepals c. 1A the length of the corolla tube or, usually, much shorter; corolla tube solid in the basal part, twice as long as the lobes or longer; outer staminodes 2, petaloid; ovary 3-locular. Fruit dehiscent; seeds arillate. Ten to fourteen species distributed from India to Indochina and in the western part of the Malesian region (Andersson 1998). Three

species in Sabah.

NOTES. No type was designated by Schumann when he described the genus (Schumann 1902; Greuter et al. 1993). As lectotype we have chosen Stachyphrynium latifolium (Blume) K. Schum. because it has the simple spike-like inflorescence that Schumann mentions in his protologue.

KEY TO THE SPECIES OF STACHYPHRYNIUM

1. Synflorescence capitate; bracts stiff and pointed ........ 5.3. S. placentarium Synflorescence spiciform, bracts not pointed ........... ............ 2

2. Inflorescence simple, borne on a separate shoot; synflorescence 12 - 16 cm long; plant up to 3 m tall; leaf blade 49 - 60 x 28 - 32 cm ...... 5.2. S. latifolium

Inflorescence branched, interfoliar; synflorescence 2 - 4 cm long; plant up to 0.8 m tall; leaf blade 15 - 22 x 4- 7 cm ................. 5.1. S. borneense

5.1. Stachyphrynium borneense Ridl., J. Bot. 75: 204 (1937). Type: Malaysia, Sarawak, base of Mount Matang, H. N. Ridley 11796 (holotype K!).

Rhizomatous ground herb, up to 80 cm tall. Leaves 2 - 4 per shoot; sheath 9 - 12 cm long, green to brown, glabrous; petiole 12 - 39 cm long, green, glabrous; pulvinus 1 - 2 cm long, green, glabrous; lamina oblong-elliptic, 15 - 22 x 4 - 7 cm, dark green above, pale green below, slightly plicate along the primary veins. Inflorescence interfoliar, erect; peduncle 5 - 10 cm long, glabrous; synflorescence 2 - 4 cm long, spiciform, with 1 - 4 branches of up to 2 orders; fertile bracts 2 - 6 per branch, brownish red, oblong to lanceolate, c. 16 x 5 mm, slightly hairy at the tip; special paraclades with 2 - 3 flower pairs, associated prophylls c. 14 x 2 mm, interphylls c. 12 x 2 mm. Flower c. 2 cm long, slender, white; sepals subulate, c. 3.8 x 0.3 mm; corolla tube c. 16 mm long, solid 6 - 7 mm above the ovary; petal lobes subulate, c. 6 x 1.5 mm; staminal tube c. 3 mm longer than the corolla tube; outer staminodes equal, obovate, petaloid lobes 4.5 x 3.8 mm; cucullate staminode 2 x 1.8 mm; callose staminode 1.5 x 1.5 mm; stamen 1.3 mm long; style free part c. 3 mm long, curved, stigmatic cavity 0.9 mm diameter; ovary c. 1.5 mm long. Fruit reddish yellow, round, oblong, c. 1.3 cm long; seeds 1- 3, 7 mm long, globose to ellipsoid, with a large, light, bifid aril, 6 mm long. Fig. 8A.


DISTRIBUTION. Widely distributed in N and W Borneo (Sabah, Sarawak and Brunei). The species may also occur in Kalimantan although we have not seen any specimens from there. Whether the species occurs outside Borneo is uncertain (see discussion of S. sumatranum in the notes below).

HABITAT AND ECOLOGY. Found in primary or secondary forest, in deep shade. Stachyphrynium borneense is capable of covering large areas of forest floor (pers. obs.). Observations of flowering phenology (Clausager unpubl.) have shown that the flowers open after dawn and only live for one day. Each synflorescence usually has one open flower or none each day, and can produce flowers for 1 to 2 months.

SELECTED SPECIMENS EXAMINED (from a total of 9). SABAH: Keningau distr., Tambulanan, Gambukas 69090 (SAN); Lahad Datu distr., Jalan Teck Guan, Takun, Ulu Bakapit, Saigol 93116 (SAN); Tawai Forest Reserve, Telupid, Sandakan, Zainudin 5005 (KEP); Tabin Wildlife Reserve, trail to Lipad mud volcano, Borchsenius 455 (AAU); Danum Valley Conservation Area, West trail, Clausager 6 (AAU); Tawau Hills National Park, trail to sulphur spring, Clausager 21 (AAU); Maliau Basin, trail from Bangbangan camp to Maliau Falls, Poulsen 1648 (AAU); secondary forest near Imbak R., Poulsen 1588 (AAU).

NOTES. Stachyphrynium borneense strongly resembles S. sumatranum (Miq.) K. Schum. According to Ridley (1937), the difference between these two species is that plants of Stachyphrynium borneense form a "densely caespitose mass", whereas S. sumatranum "appears to have a long rhizome with distant leaf-tufts". According to our field observations S. borneense can develop both distant leaf-tufts and more dense clusters of shoots. We have not been able to examine the type of Stachyphrynium sumatranum (Teysmann s.n., BO?), and therefore we prefer not to formally synonymise the two names; if they are conspecific then sumatranum is the older epithet (1860).

A specimen collected in Sandakan in 1924 (Kloss 18734, SING!), identified as Stachyphrynium cf. jagorianum is S. borneense.

5.2. Stachyphrynium latifolium (Blume) K. Schum. in Engler, Das Pflanzenreich IV, 48: 49 (1902).

Phrynium latifolium Blume, Enum. P1. Javae: 37 (1827). Type: Indonesia, Java, Bantam province, Blume s.n. (holotype L!).

Phrynium griffithii Baker in Hook. f., Fl. Brit. India VI: 260 (1892); synon. nov. Type: Malacca, Griffith s.n. (K!)

Stachyphrynium griffithii (Baker) K. Schum. in Engler, Das Pflanzenreich IV, 48: 49 (1902).

Rhizomatous ground herb up to 3 m tall. Leaves 1 - 3 per shoot; sheath 30 - 48 cm long, yellow-brown, glabrous; petiole 140 - 180 cm long, green, glabrous; pulvinus 8 - 11 cm long, green, glabrous; lamina ovate, 49 - 60 x 28 - 32 cm, glabrous, upper surface green, with a lighter channelled midrib, lower surface light green with a raised, yellow-green midrib. Inflorescence borne directly from the rhizome, erect, simple, usually positioned close to the vegetative shoot; peduncle 17 - 32 cm long, green, glabrous; main inflorescence axis 12 - 16 cm long; fertile


4 ??ZVI A

NO J-1 .? 1p;

AMC

R 11.

. t?u

AY . . . . . . . . . .

FIG. 8. A Stachyphrynium borneense, synflorescences. Photo F. Borchsenius; B Stachyphrynium latifolium, close up of

synflorescence. Photo K. Clausager.

bracts 13 - 16, distichous, green when young, becoming brown with age, glabrous, ovate, 3 - 4.5 x 2 - 3 cm, spreading and becoming flattened with age; flower pairs 3 - 4 per special paraclade, associated prophylls 2 - 3 x 1 - 1.5 cm, interphylls 1.5 -

2.5 x 0.5 - 1 cm. Flower c. 4 cm long, white, very slender; sepals subulate, 5.4 x 1.5 mm; corolla tube 25 mm long and 1.2 mm, the basal 6 mm solid; petal lobes subulate, 11 x 6 mm; staminal tube of same length as the corolla tube; outer staminodes sub-equal, petaloid lobes 9 - 11 x 6- 9 mm; cucullate staminode 5 x 3 mm, yellow at the tip; callose staminode 6 x 4 mm; fertile stamen 3 mm long, with a

petaloid appendage of 4.5 x 3 mm; style with free part 6 mm long, curved, stigmatic cavity 1.2 mm diameter; ovary 4 mm long. Fruit (description taken from Holttum 1951) 2.3 cm long, with 2 - 3 rugose seeds. Fig. 8B.

DISTRIBUTION. Widespread in the west Malesian region, from S Thailand and the Malay Peninsula to Java, Sumatra and Borneo. In Sabah, S. latifolium has been found only east of the Crocker Range.

HABITAT AND ECOLOGY. Some of the specimens examined were specifically noted as growing on limestone, but the species is not limited to this habitat. Our own collections were made in disturbed forest, or from cultivated plants.

SELECTED SPECIMENS EXAMINED (from a total of 16). SABAH: Kinabalu National

Park, along Sng, Mamut, near Poring, Kokawa & Hotta 4887 (L); Kinabalu National Park, along trail between Poring and Langana water fall, Tamura & Hotta 233 (L); Kinabalu National Park, Poring Hot Springs, trail from Canopy walkway, Clausager 16 (AAU, SAN); Lamag distr., Batu Puteh, Majawat & Lasan 88013 (SAN); Km 29, Jalan Sukau, Bukit Garam, Kinabatangan, George et al. 117680 (SAN); Tenom Agricultural Station, Native Orchid Centre (cultivated), Clausager 7 (AAU, SAN).


NOTES. The type specimens of Stachyphrynium latifolium and S. griffithii show no differential characters and hence we have synonymised the two names. Another species of the same group, S. cylindricum (Ridl.) K. Schum., distributed in southern Thailand and the northernmost part of the Malay Peninsula, is very similar to S. latifolium, but has more narrow inflorescences and may represent a distinct species. Further studies are needed to clarify this relationship.

5.3. Stachyphrynium placentarium (Lour) Clausager & Borchs. comb. nov.

Phyllodes placentaria Lour., Fl. Cochinch.: 13 (1790). Type: plate in Rumphius Herbarium Amboniense 1. II. c. 12. t. 53 (1741).

Phrynium placentarium (Lour.) Merr., Philipp.J. Sci. 15: 230 (1919). Phrynium parviflorum Roxb., Hort. Bengal.: 1 (1814); Fl. Ind. 1: 7 (1820); Baker in

Hooker f., Fl. Brit. India VI: 259 (1892); K. Schumann in Engler, Das Pflanzenreich IV, 48: 54 (1902). Type: Wallich, Catalogue 6620? (K-W, microfiche at AAU!).

Phrynium densiflorum Blume, Enum. Pl. Javae: 38 (1827). Type: Indonesia, Java, Blume s.n. (lectotype plus isotype L!).

Rhizomatous ground herb up to 1.5 m tall. Leaves 1 - 3 per shoot; sheath 32 - 37 cm long, green, covered by whitish, short hairs; petiole 37 - 47 cm long, green, glabrous; pulvinus 3 - 5 cm, light green to yellow, glabrous; lamina 31 - 46 x 12 - 15 cm, green above, light green beneath, glabrous. Inflorescence interfoliar, erect, sometimes appearing to protrude from the petiole of the accompanying leaf; peduncle 77 - 93 cm long, green, glabrous; synflorescence capitate, 6 - 8 cm diameter, made up by 5 - 10 densely congested branches, 3 - 4 cm long; bracts subtending the proximal branches enlarged, 3 - 5.5 cm long, stiff, acuminate, light green and slightly hairy; fertile bracts spirally or

distichously arranged, light green with cream or white margin, 2.5 - 3 x 1.5 - 2 cm, lanceolate, stiff and sharply pointed at apex; flower pairs 2 - 3 per special paraclade, associated prophylls c. 2 x 0.7 cm, interphylls 1.5 - 2 x 1 - 1.5 cm. Flowers white, c. 2.7 cm long; sepals subulate, 3.6 x 0.3 mm; corolla tube 18 mm

long; petal lobes subulate, 6 x 3 mm; staminal tube c. 3 mm longer than the corolla tube; outer staminodes slightly unequal, petaloid lobes 3 - 4.5 x 2.3 mm; cucullate staminode 3 x 1.5 mm; callose staminode 2.3 x 1.8 mm; stamen 2.2 mm

long; style with free part c. 3 mm long, curved, stigmatic cavity 0.6 mm diameter; ovary 1.8 mm long. Fruit blue-grey, 1.5 - 2 cm in diameter, globose; seeds 1 - 3, blue, with a red to pink, bifid aril.

DISTRIBUTION. Widespread in Indochina and S China, Thailand, Peninsular Malaysia, Sumatra, Java, and Borneo. In Sabah, Stachyphrynium placentarium has been found in a narrow belt on the eastern side of the Crocker Range, from Serinsim in the north, through Ranau, to Tenom in the south.

HABITAT AND ECOLOGY. Stachyphrynium placentarium is commonly found in

moderately disturbed vegetation, often together with Phrynium pubinerve. SPECIMENS EXAMINED. SABAH: Tenom Valley, rubber plantation behind Agricultural


Station, Clausager 12 (AAU); Tenom Distr., Melurut Forest Reserve, Ulu Sg. Oloson Melalap, Mantor 96329 (SAN); along Sungai Bambangan, N of Ranau, alt. 800 - 850 m, Tamura & Hotta 446 (L); Ranau Distr., Sg. Bambangan, Amin et al. 123575 (SAN); Kota Marudu Distr., Kampung Serinsim, Bakia 73 (Sabah Parks); Kota Marudu Distr., Kampung Serinsim, Bakia 233 (Sabah Parks).

NOTES. This species was placed in the genus Phrynium by Merrill (1919) because of its capitate synflorescence. However, the flowers correspond to those of Stachyphrynium in having very short sepals and a very long, slender corolla tube.

INCOMPLETELY KNOWN SPECIES

Tarmiji SAN 82864, (SAN), from the Sandakan District, appears to be a Phacelophrynium species. This specimen has a synflorescence that is extremely elongate and relatively sparsely branched compared to other species in this genus. A similar collection from the Ranau district (Amin & David SAN 109768) has been determined as Phrynium capitatum, but this is clearly incorrect. The leaves of both specimens show no signs of variegation, and in both cases the flowers are noted as being white. Unfortunately we have not seen this species in the field, and there is no pickled flower material accompanying the two collections. It has therefore not been possible to make a proper description, nor to determine unambiguously which genus the species belongs to. A third collection, apparently representing the same species, also from the Sandakan district (S. Lantoh SAN 87908), includes an infructescence; here the flowers are noted as being yellow-white, and the fruits as being yellow-brown.

ACKNOWLEDGEMENTS

The study was carried out as a part of the project 'Collaboration on Biodiversity between Universiti Malaysia Sabah and Danish Universities', funded by Danced (Danish Co-operation on Environment and Development). We gratefully acknowledge the help from, and collaboration with, the following people and institutions: Dr Maryati Mohamad and Mr Berhaman Ahmad of the UMS; Yayasan Sabah; the Royal Society; Danum Valley Field Centre; Sabah Parks; and the Forest Research Centre in Sepilok. Anthony Lamb shared his extensive knowledge of Sabah Marantaceae with us and pointed out several field localities; we are extremely grateful for this assistance. Helen Kennedy and John Mood are thanked for inspiring comments and for drawing our attention to undescribed species. Curators of the following herbaria are thanked for allowing us to access their collections: A, BM, C, K, KEP, L, P, SAN, SING, UKM and Sabah Parks Herbarium. The authors are indebted to Benjamin 0llgaard for assistance with the Latin diagnoses and for his help with nomenclatural puzzles. Jens Christian Schou made the line drawings. Finally, we thank Dr Axel

Dalberg Poulsen for his help and support during field work, constructive comments, and permission to use his photos as illustrations.


REFERENCES

Andersson, L. (1976). The synflorescence of the Marantaceae. Organization and

descriptive terminology. Bot. Not. 129: 39 - 48. (1977). The genus Ischnosiphon (Marantaceae). Opera Bot. 43: 1 - 114. (1981). The neotropical genera of Marantaceae. Circumscription and

relationships. NordicJ. Bot. 1: 218 - 245. - (1986). Revision of Maranta subgen. Maranta (Marantaceae). Nordic J. Bot. 6

(6): 729 - 756. - (1998). Marantaceae. In: K. Kubitzki (ed.), The Families and Genera of

Vascular Plants, pp. 278 - 293. Springer Verlag, Berlin. - & Chase, M. W. (2001). Phylogeny and classification of Marantaceae. Bot. J.

Linn. Soc. 135: 275 - 287. Blume, C. L. (1827). Enumeratio Plantarum Javae, pp. 37 - 38. J. W. van Leeuwen,

Leiden. Christensen, H. (1997). Uses of plants in two indigenous communities in Sarawak,

Malaysia. Part of Ph.D thesis, University of Aarhus, Denmark. Greuter, W., Brummitt, R. K., Farr, E., Kilian, N., Kirk, P. M. & Silva, P. C. (1993).

Names in Current Use for Extant Plant Genera. Koeltz Scientific Books. Germany. - , McNeill, J., Barrie, F. R., Burdet, H. M., Demoulin, V., Filgueiras, T. S.,

Nicolson, D. H., Silva, P. C., Skog, J. E., Trehane, P., Turland, N.J. & Hawksworth, D. L. (eds.) (2000). International Code of Botanical Nomenclature (Saint Louis Code). Regnum Vegetabile 138. Koeltz Scientific Books, Germany.

Holmgren, P. K., Holmgren, N. H. & Barnett, L. C. (1990). Index Herbariorum, eighth edition. New York Botanical Garden, New York.

Holttum, R. E. (1951). The Marantaceae of Malaya. Gard. Bull. Singapore 13: 254 - 269. Kato, M. (1996). Plant-pollinator interactions in the understory of a lowland mixed

dipterocarp forest in Sarawak. Amer. J. Bot. 83: 732 - 743. Kennedy, H. (1978). Systematics and pollination of the "closed flowered" species of

Calathea (Marantaceae). Univ. Calif. Publ. Bot. 71: 1 - 90. (1986). Two new striped-leaved species of Calathea (Marantaceae) from

Ecuador. Canad. J. Bot. 64: 1321 - 1326. Kirchoff, B. (1986). Inflorescence structure and development in the Zingiberales:

Thalia geniculata (Marantaceae). Canad. J. Bot. 64: 859 - 864. Kress, W. J. (1990). The phylogeny and classification of the Zingiberales. Ann.

Missouri Bot. Gard. 77: 698 - 721. Kunze, H. (1983). Typological interpretation of the inflorescence of the

Marantaceae. Acta Bot. Neerl. 32: 370. - (1985). Die Infloreszenzen der Marantaceen und ihr Zusammenhang mit

dem Typus der Zingiberales-Synfloreszenz. Beitr. Biol. Pflanzen 60: 93 - 140. - (1989). Probleme der Infloreszenztypologie von W. Troll. P1. Syst. Evol. 163:

187 - 199. Linnaeus, C. (1753). Species Plantarum. Stockholm. Merrill, E. D. (1919). Phrynium. Philipp. J. Sci. 15: 230.

- (1924). Marantaceae. In: An enumeration of Philippine flowering plants 6: 248 - 251. Bureau of Printing, Manila.


Momose, K., Yomoto, T., Magasitu, T., Kato, M., Nagamasu, H., Sakai, S., Harrison, R. D., Itioka, T., Hamid, A. A. & Inoue, T. (1998). Pollination biology in a lowland

dipterocarp forest in Sarawak, Malaysia. I. Characteristics of the plant pollinator community in a lowland dipterocarp forest. Amer. J. Bot. 85: 1477 - 1501.

Ridley, H. N. (1899). The Scitaminee of the Malay Peninsula. J. Straits Branch Roy. Asiat. Soc. 32: 175 - 182.

- (1906). Scitaminee of Borneo. J. Straits Branch Roy. Asiat. Soc. 46: 229 - 246. S (1909). The Scitaminec of the Philippine Islands. Philipp. J. Sci., C 45: 155 - 199. - (1937). New Asiatic Scitamineae. J. Bot. 75: 204 - 205.

Rolfe, R. A. (1907). Donax and Schumannianthus. J. Bot. 45: 242 - 244. Schumann, K. (1893). Marantaceae africanae. Bot. Jahrb. Syst. 15: 428 - 446.

S(1902). Marantaceae. In: A. Engler (ed.), Das Pflanzenreich IV. 48, pp. 1 - 148.

Verlag von Wilhelm Engelmann, Leipzig. Suresh, C. R. & Nicolson, D. H. (1986). Two nomenclatural novelties based on

Rheede's "Hortus Malabaricus". Taxon 35: 354 - 355. Turner, I. M. (1994). Notes on the flora of Malaya: New records, overlooked records

and some nomenclatural clarification. Gard. Bull. Singapore 46: 127 - 129. (2000). The Plant Taxa of H. N. Ridley, 3. The Zingiberales. Asian J. Trop. Biol.

4:1 - 47.

LIST OF EXSICCATAE

The specimens are arranged alphabetically according to the name of the collector. Determination is indicated in brackets using the abbreviations given below. Herbarium acronyms are according to Holmgren et al. (1990).

Don.can Donax canniformis (G. Forst.) K. Schum. Pha.aur Phacelophrynium aurantium Clausager & Borchs. Pha.lax Phacelophrynium laxum Clausager & Borchs. Pha.max Phacelophrynium maximum (Blume) K. Schum.

Phr.gra Phrynium grandibracteatum Clausager & Borchs. Phr.hir Phrynium hirtum Ridl.

Phr.pub Phrynium pubinerve Blume Phr.ste Phrynium stenophyllum Clausager & Borchs. Phr.vil Phrynium villosulum Miq. Sch.dic Schumannianthus dichotomus (Roxb.) Gagnep. Sta.bor Stachyphrynium borneense Ridl. Sta.lat Stachyphrynium latifolium (Blume) K. Schum.

Sta.pla Stachyphrynium placentarium (Lour.) Clausager & Borchs.

Aban 83169 (Pha.max) L; Aban & Petrus 90661 (Pha.max) K; Ahmad 1100 (Phr.pub) SING; Alvins, V M. 2277 (Phr.hir) SING; Ambri 1580 (Phr.gra) SAN; 1577 (Phr.gra) K, SAN; Amdjah 145 (Pha.lax) L; 430 (Phr.pub) L; Amin 111544 (Don.can) SAN; 123575 (Sta.pla) SAN; Amin &Jarius 114279 (Pha.max) KEP; Amin & Donggop 93943 (Pha.max) L; Amin & Good 100225 (Phr.pub) SAN; Amin et al. 111544


(Don.can) SAN; 123575 (Sta.pla) SAN; Andrews, S. 729 (Pha.aur) UKM; Awa, D. & Yii, P.C. 46718 (Pha.aur) K; 46664 (Phr.max) K.

Bakia, K. 73 (Sta.pla) Sabah Parks; 233 (Sta.pla) Sabah Parks; Beaman, J. H. 10606 (Pha.max) A; Beccari, 0. 630 (Pha.max) K; Best, G. A. 14110 (Phr.pub) SING; Blewett, J B. J. 3 (Pha.aur) UKM; Blume s.n. (Pha.max) L; s.n. (Phr.pub) L; s.n. (Sta.lat) L; Borchsenius, E 455 (Sta.bor) AAU; 531 (Pha.aur) AAU; Burley, J.S. & Tukirin 1428 (Pha.max) SING.

Castro 3195 (Don.can) A; Chai, P 36205 (Pha.aur) KEP; Christensen, H. 1663 (Pha.max) AAU; Christensen, H. & Apu, F L. 56 (Pha.max) AAU; Christensen, H. & Poulsen, A. D. 1998 (Phr.ste) AAU; Clausager, K. 6 (Sta.bor) AAU, SAN; 7 (Sta.lat) AAU, SAN; 10 (Phr.vil) AAU, SAN; 11 (Pha.aur) AAU, SAN; 12 (Sta.pla) AAU, SAN; 13 (Phr.vil) AAU, SAN; 14 (Phr.ste) AAU, SAN; 16 (Sta.lat) AAU, SAN; 17 (Phr.vil) AAU, SAN; 18 (Phr.hir) AAU, SAN; 19 (Pha.aur) AAU, SAN; 20 (Phr.gra) AAU, SAN; 21 (Sta.bor) AAU, SAN; 22 (Pha.aur) AAU, SAN; Clausager K. & Borchsenius, F 01 (Pha.lax) AAU, SAN; 2 (Phr.hir) AAU, SAN; 3 (Phr.pub) AAU, SAN; 4 (Pha.max) AAU, SAN; 5 (Phr.pub) AAU, SAN; Clausager, K. & Poulsen, A. D. 8 (Phr.max) AAU, SAN; Clemens, J. & Clemens, M. S. 22137 (Pha.aur) K; 27204 (Pha.max) K; Corner, E. J. H. 30292 (Pha.max) K; 30957 (Phr.vil) SING; 30680 (Phr.vil) SING; 30265 (Phr.hir) K; 30769 (Phr.pub) SING; 31635 (Phr.pub) SING; s.n. (Phr.pub) SING; Cowley, J. 56 (Pha.max) AAU; 181 (Pha.max) AAU; Cuadra, A. 1262 (Don.can) A; 1462 (Don.can) SAN.

Dewol et al. 113678 (Phr.pub) SAN; Dolois, S. 09001 (Pha.aur) Sabah Parks.

Edano 11905 (Pha.max) BM; Elmer A. D. E. 20255 (Don.can) A; 20945 (Phr.vil) L; 16145 (Pha.max) BM.

Fidilis 125289 (Pha.aur) KEP; Fidilis & Sumbing 68784 (Don.can) SAN; 68787

(Phr.pub) SAN; 95701 (Phr.pub) SAN; Forbes, H. 0. 2987 (Sta.lat) BM; 511 (Pha.max) BM; Forman, L. L. & Blewett, J. B. J. 1200 (Sta.bor) SING; Forster s.n. (Don.can) BM.

Gambukas, S. 69090 (Sta.bor) SAN; George et al. 117680 (Sta.lat) SAN; Gibot, A. 32968 (Don.can) SAN; 66805 (Don.can) SAN; 97066 (Pha.max) L; Goodenough, J. S. 4610 (Sta.lat) SING; s.n. (Sta.lat) SING.

Hansen, C. 241 (Pha.lax) C; 619 (Pha.lax) C, KEP, SAN; 920 (Pha.lax) C; Haviland, G. D. s.n. (Phr.vil) K; s.n. (Phr.vil) K; Henderson, M. R. 21850 (Phr.pub) SING; 22533

(Phr.pub) SING; 10397 (Phr.pub) SING; Holttum, R. E. 19858 (Sta.lat) SING; 9526

(Phr.pub) SING.

Jimpin, S. 106981 (Pha.aur) K; Johannsen, S. 9 (Pha.max) AAU, SAN; 11 (Phr.gra) AAU, SAN; 13 (Phr.vil) AAU, SAN;Joseph, B. 123696 (Pha.max) KEP;Joseph, S. 56759 (Don.can) SAN.

Kadir 2054 (Pha.aur) A; Kadis 2575 (Don.can) A; Kennedy, H. & Ping, T. E. 4464


(Sta.lat) SING; Kiah 35410 (Sta.lat) SING; 13337 (Pha.max) SING; Kloss & Robinson 6956 (Sta.lat) K; Kokawa, S. & Hotta, M. 2443 (Phr.vil) SAN; 2625 (Don.can) L; 4843 (Phr.pub) L; 4887 (Sta.lat) L; 4963 (Phr.pub) L; Krispinus, E 95643 (Don.can) SAN; 95447 (Don.can) SAN; 91920 (Don.can) SAN.

Lantoh, M. 109675 (Pha.aur) KEP; Lassan, P 116818 (Phr.pub) SAN; Lee, Y F & Aban, G. 96785 (Don.can) SAN; 124807 (Pha.aur) SAN; Lideh, J. B. & Donggop 116974 (Pha.max) KEP; Loureiro s.n. (Don.can) BM.

Madani, L. 112946 (Pha.aur) KEP; 90839 (Pha.max) L; 132698 (Phr.hir) SAN; Maidin 2677 (Don.can) A; Maikin & Lidah 131127 (Pha.aur) SAN; Majawat, G. & Lassan, P 88013 (Sta.lat) SAN; Mansus et al. 108856 (Don.can) SAN; Mansus & Dewol 109237 (Phr.pub) SAN; Mantor, A. 96329 (Sta.pla) SAN; Meijer, W 131944 (Pha.aur) A; 05611 (Pha.aur) Sabah Parks; Merrill 7288 (Pha.max) BM.

Ngadiman 37084 (Pha.max) BM, SING; Nooteboom, H. P 1240 (Don.can) L; 1365 (Phr.pub) L;

Nut, M. 18554 (Phr.pub) SING.

Ogata, K. 10896 (Don.can) L; Orolfo 55486 (Pha.max) KEP; Othman, H. 61748 (Pha.max) K.

Pikkoh, M. 101443 (Don.can) SAN; Poulsen, A. D. 1588 (Sta.bor) AAU; 1602 (Sta.bor) AAU; 1648 (Sta.bor) AAU; 1656 (Phr.vil) AAU; Poulsen, A. D. & Abdilla, R. 1683 (Pha.max) AAU; Poulsen, A. D. & Banggilon, L. 1650 (Pha.max) AAU; Poulsen, A. D. & Borchsenius, E 1576 (Pha.max) AAU; 1575 (Pha.hir) AAU; 1574 (Phr.pub) AAU; Purseglove, J W 5302 (Pha.aur) K

Ramos 15229 (Pha.max) BM; 1488 (Phr.hir) K; Rena et al. 58360 (Pha.max) K; Rhea, G. & Aban 76834 (Don.can) SAN; Ridley, H. N. 44 (Sta.lat) SING; 444 (Sta.lat) SING; 2398 (Pha.max) K; 9787 (Sta.lat) K; 10000 (Phr.vil) K; 11796 (Sta.bor) K; 13272 (Phr.hir) K; 14036 (Phr.vil) K, SING; s.n. (Phr.hir) SING; s.n. (Phr.pub) SING; s.n. (Sta.lat) SING.

Saigol, P 93114 (Phr.pub) SAN; 93116 (Sta.bor) SAN; Sam, P P 49917 (Phr.pub) SAN; Shah, M. 1629 (Phr.pub) SING; 4970 (Phr.pub) SING; Shah, M. &

Shukor, A.

2322 (Pha.max) SING; Sinclair, J. 40040 (Phr.vil) SING; Sinanggol, H. T 57434 (Don.can) SAN; Stevens,

P.E 399 (Phr.pub) SAN; 499 (Pha.aur) KEP; 649 (Pha.max)

L; Stone, B. C. 85189 (Pha.aur) L, SAN; Sundaling, D. 80772 (Don.can) SAN; 83918 (Don.can) SAN; 91084 (Phr.pub) SAN; 96992 (Pha.max) L.

Tamura, M. & Hotta, M. 233 (Sta.lat) L; 446 (Sta.pla) L; 879 (Don.can) L; Tandom 2833 (Sch.dic) L; Teijsmann 4490 (Phr.vil) U.

Vidal 3976 (Pha.max) K.

Wenzel 232 (Pha.max) BM.

Zainudin, A. 4822 (Pha.max) UKM; 5005 (Sta.bor) UKM.


INDEX OF SCIENTIFIC NAMES

Names of recognized taxa are in bold.

Actoplanes K. Schum.

canniformis (G. Forst.) K. Schum. Donax Lour

arundastrum Lour. canniformis (G. Forst.) K Schum. grandis (Miq.) Ridl.

Heliconia L.

triumphans Linden ex Rodigas Maranta L.

grandis Miq. Phacelophrynium K Schum.

aurantium sp. nov. bracteosum K. Schum. laxum sp. nov. maximum (Blume) K Schum.

tapirorum (Ridl.) K. Schum. Phrynium Willd.

basiflorum Ridl. basiflorum var. nobile Ridl. capitatum Willd. densiflorum Blume

grandibracteatum sp. nov.

griffithii Baker hirtum Ridl. inflatum Merr.

latifolium Blume malaccense Ridl. maximum Blume

parviflorum Roxb.

placentarium (Lour.) Merr.

pubinerve Blume rheedei Suresh & Nicolson

stenophyllum sp. nov.

tapirorum Ridl. villosulum Miq.

Phyllodes Lour.

placentaria Lour.

Pontederia L. ovata L.

Schumannianthus Gagnep. dichotomus (Roxb.) Gagnep.

Stachyphrynium K Schum. borneense Ridl.

cylindricum (Ridl.) K Schum.

griffithii (Baker) K. Schum. latifolium (Blume) K Schum.

placentarium (Lour) Clausager & Borchs. comb. nov.

sumatranum (Miq.) K Schum.

Documents

The Marantaceae of Sabah, Northern Borneo