A monograph of the genus Orania (Arecaceae: Oranieae)

Summary

Orania Zipp. (Oranieae: Arecoideae: Arecaceae) is a genus of solitary, single stemmed tree palms consisting of 28 species, of which 11 species are new and described for the first time in this paper. The genus has an interesting disjunct distribution, with 25 species found in Malesia and three species confined to Madagascar. The 11 new species, O. bakeri A. P. Keim & J. Dransf., O. dafonsoroensis A. P. Keim & J. Dransf., O. deflexa A. P. Keim & J. Dransf., O. ferruginea A. P. Keim & J. Dransf., O. grandiflora A. P. Keim & J. Dransf., O. littoralis A. P. Keim & J. Dransf., O. longistaminodia A. P. Keim & J. Dransf., O. subdisticha A. P. Keim & J. Dransf., O. tabubilensis A. P. Keim & J. Dransf., O. timikae A. P. Keim & J. Dransf. and O. zonae A. P. Keim & J. Dransf. are found in New Guinea. A determination key is produced and detailed descriptions provided for all taxa.

Introduction

The subject of this monograph is Orania Zipp., a genus of single stemmed pinnate leaved monoecious tree palms with 28 species. The genus has an interesting disjunct distribution, with 25 species in Malesia and three species confined to Madagascar. There has been no recent monograph since that of Essig (1980), yet many new collections have been made and thus the genus now needs substantial revision. With the presence of triads and reduplicate pinnate leaves, Orania is unmistakably a member of the subfamily Arecoideae (Dransfield et al. 2008).

Relationships of Orania

Drude (1887) included Orania in his subfamily Ceroxylinae. Orania together with Arenga Labill., Caryota L. and Wallichia Roxb. were included in the subtribe Caryoteae [sic.]. However, Drude did not mention the obvious difference in leaflet plication among the genera he included in the same subtribe, reduplicate in Orania and induplicate in the others. Beccari & Pichi Sermolli (1955) were the first to propose the tribe Oranieae Becc., but in their concept the tribe also included Sclerosperma G. Mann & H. Wendl. and Ravenea H. Wendl. ex C. D. Bouché. Based on pollen data, Sowunmi (1972) excluded the two genera from Oranieae and suggested separate tribes for each of them. Sowunmi suggested the tribe should be monotypic. There was no mention of Sindroa Jum., and Halmoorea J. Dransf. & N. W. Uhl had not yet been proposed.

In the first edition of Genera Palmarum (GP1), Uhl & Dransfield (1987) placed Orania (including Sindroa) together with Halmoorea in subtribe Oraniinae J. Dransf. & N. W. Uhl within the tribe Areceae. Later Dransfield & Beentje (1995) included the genus Halmoorea in Orania thus leaving Orania as the only genus in the subtribe. Uhl & Dransfield mentioned that the inclusion of Oraniinae together with the other three subtribes with triovulate-triloculate gynoecia (Manicariinae J. Dransf. & N. W. Uhl, Leopoldiniinae J. Dransf. & N. W. Uhl and Malortieinae Benth. & Hook. f.) in Areceae was problematic. However, at the time there seemed to be no better placement for these anomalous taxa (Uhl & Dransfield 1999). Henderson (1990) suggested two possible affinities for the then subtribe Oraniinae, either with the subtribe Wettiniinae Benth. & Hook. f. (tribe Iriarteeae Drude) or the monogeneric tribe Podococceae J. Dransf. & N. W. Uhl.

Uhl et al. (1995) were the first after GP1 to indicate that the genus might not be a member of tribe Areceae. The result of their study shows that Orania is in the same clade as Podococcus G. Mann & H. Wendl., which according to GP1 is not a member of Areceae. Orania is separated from the other members of Areceae except for Manicaria Gaertn. Manicaria is a member of one of the problematic subtribes within Areceae, Manicariinae (Uhl & Dransfield 1987).

In the most recent phylogenetic classification of the palm family based on molecular and morphological data, Orania is treated as the sole member of the tribe Oranieae (Dransfield et al. 2005, 2008). Unlike the classification of Beccari & Pichi Sermolli (1955), in this recent classification the genera Sclerosperma and Ravenea H. Wendl. ex C. D. Bouché are excluded from Oranieae. Sclerosperma is placed in its own tribe, Sclerospermeae, whereas Ravenea belongs in tribe Ceroxyleae within a completely different subfamily (Ceroxyloideae).

History of the genus Orania Zipp.

The first use of the generic name Orania was in a letter by Zippelius dated 1823 to Blume (Zippelius 1828a) which was later published in Algemeen Konst- en Letter-Bode 1: 297 in 1829. Zippelius also sent the same letter to Tausch in Germany (Zippelius 1828b) where it was later included in Flora oder Botanische Zeitung Regensburg (Flora) 12, also published in 1829. However, Blume’s publication has priority. Zippelius saw the first species from the new genus in Triton Bay (Papua) and named it Orania regalis. The generic name Orania itself commemorates F. G. L. Willem van Nassau, Prince of Orange and Crown Prince of the Netherlands (1792 – 1849) (Backer 1936). A picture of O. regalis (as Arausiaca excelsa, picture number 122) published by Blume (1836), which according to him was drawn based on Zippelius’s original field notes (now lost), is selected as the type.

Arausiaca Blume (1836, pictures 119 & 112), comprising one species only, A. excelsa, in New Guinea and Macrocladus Griff. (Griffith 1845), also comprising one species only, M. sylvicola, in Western Malesia (Malay Peninsula to Sumatra and Java), were described at about the same time. However, in 1849 Carl von Martius included Macrocladus Griff. in Orania and Miquel (1859) included Arausiaca Blume in Orania.

Beccari suggested Orania beccarii F. M. Bailey indicated the presence of Orania in Australia based on specimens sent to him by Bailey (1909), noting, however, several characters not shared with the rest of Orania and he thus named a distinct subgenus, Oraniopsis, with O. beccarii as the only member (Beccari & Pichi-Sermolli 1955). Later this species was treated as a synonym of Orania appendiculata (F. M. Bailey) Domin (Domin 1915). O. appendiculata was later excluded from Orania and put in the completely unrelated genus Oraniopsis (Dransfield et al. 1985). Based on its morphological characters this species belongs to the completely different subfamily, Ceroxyloideae. This ended the 75 year division of Orania into two subgenera and also showed that there is no Orania living in the wild in present day Australia.

Although Beccari is known to have conducted several botanical explorations in East Malesia, including Moluccas and New Guinea (Beccari 1877, 1905, 1915; Van Steenis 1950), he did not make as many collections of Orania there as botanists prior to him did from West Malesia, kept in BO and FI. This is reflected in his more detailed descriptions of taxa (both species and varieties) that occur in West Malesia compared to Moluccas and New Guinea. Beccari described only four taxa of Orania from East Malesia (O. aruensis, O. lauterbachiana, O. micrantha and O. moluccana) compared to seven in West Malesia. The other New Guinean taxa known to Beccari (O. macropetala K. Schum. & Lauterb. and O. regalis) had already been described previously by other botanists.

Burret described all of his species of Orania (O. archboldiana, O. brassii, O. clemensiae and O. disticha) based on herbarium specimens sent to him by other botanists who worked in New Guinea, such as L. J. Brass, M. S. Clemens, K. A. Lauterbach and K. Schumann. Burret had more New Guinean specimens available to him and this is reflected in the greater number of species from New Guinea he described (four) compared to Beccari (two). However, Burret is notorious for his narrow species concept.

Jumelle (1933) published the genus Sindroa with one species. Thanikaimoni (1970) grouped Orania and Sindroa together based on the similarity of their pollen type (which is monosulcate with tectate exine) into his first of 27 pollen types. However, he still recognised them as two distinct genera. Sindroa was later shown by Dransfield & Uhl (1984) to be synonymous with Orania. In the same publication Dransfield & Uhl also described a new genus related to Orania found in Madagascar, Halmoorea, with only one species H. trispatha. Dransfield & Beentje (1995) later included Halmoorea as a synonym of Orania based on much more extensive material.

The last synopsis of the genus prior to this study was published by Essig in 1980; however, in this synopsis Essig still mentioned only 16 species, 15 in Malesia and one in Australia (Orania appendiculata). He described four new species from Papua New Guinea. Three species (O. brassii, O. clemensiae and O. micrantha respectively) were lumped into O. lauterbachiana. The presence of Orania in Madagascar had not been recognised at the time of Essig’s study. Essig conducted more botanical explorations than either Beccari or Burret, particularly in the eastern part of New Guinea (Papua New Guinea). This is reflected in the increased number of species in his synopsis of the genus. Some of the New Guinean taxa described by Beccari and Burret were placed by Essig as synonyms of other species, based on his assessments of the level of morphological difference. Unfortunately, Essig was not able to explore in the western half of the island (Papua), because the province was closed for foreign scientific exploration for most of the 1980s. Therefore, apart from O. brassii, O. clemensiae and O. micrantha, Essig still followed Burret for the other Papuan taxa. Essig did not conduct any botanical exploration in the other parts of Malesia and merely followed Beccari when treating the extra-New Guinean taxa.

The most recent list of Orania in New Guinea was published by Ferrero (1997). He followed the synopsis by Essig; however, he worked only in mainland New Guinea. Although he was able to work in Papua and Papua New Guinea, the number of species recognised by him agreed with Essig (1980), apart from one extra taxon from the Toricelli Mountains in the northern part of the border between Papua and Papua New Guinea with distichous leaves proposed as a new species but not formally described.

In our synopsis the increased number of species is due partly to more extensive botanical explorations, especially in New Guinea and Madagascar and due also to a different opinion concerning several morphological characters considered unimportant by Essig. This affects particularly the treatment of Orania lauterbachiana. However, Essig’s four new species are still accepted.

Table 1 shows the number of taxa (including species and varieties) of Orania recognised in the current study compared to the treatments of Beccari (1877 – 1919), Burret (1935 – 1940), Essig (1980) and Ferrero (1997).

Table 1 Species of Orania recognised in this monograph (including their distributions) compared with previous treatments.

Distribution, ecology and uses of the genus Orania

Twenty-five of the 28 species of Orania are found within Malesia. Only one of those species, O. sylvicola (Griff.) H. E. Moore, penetrates into a small, restricted area beyond the border with the Indochinese floristic region (southern part of Thailand). The other three species are distributed in Madagascar (Baker et al. 1998). In Malesia Orania is found in almost every part of the floristic region from the southern part of Thailand to the Malay Peninsula, Sumatra, Java (western part only), Borneo, the Philippines, possibly in Sulawesi (Celebes), the Moluccas and mainland New Guinea including the small nearby islands offshore in Milne Bay in the south-east (D’ Entrecasteaux Islands). The centre of diversity is in New Guinea, both in Papua and Papua New Guinea, where 22 species have been recognised in this study. Nineteen of those 22 species are endemic to mainland New Guinea. The three other species (O. lauterbachiana, O. palindan Merr. and O. regalis) have extra-New Guinean distributions.

Orania palindan is known to have been collected or seen in the Moluccas, Celebes (Beckwith 1940 pers. not. in Fairchild 1943) and the Philippines, so is the most widespread species in the genus. O. lauterbachiana has been collected from the D’ Entrecasteaux Islands as well as from mainland New Guinea. O. regalis has also been collected in the Aru Archipelago. The genus has never been collected east of mainland New Guinea and the D’ Entrecasteaux Islands.

In Madagascar Orania is known from forest areas in the eastern and north-eastern areas and a small enclave in the north-western area. Three species have been recognised: O. longisquama (Jum.) J. Dransf. & N. W. Uhl, O. ravaka Beentje and O. trispatha (J. Dransf. & N. W. Uhl) Beentje & J. Dransf. O. longisquama is widespread, whereas the two other species are found only in the eastern and north-eastern areas. The only species reported from the islands surrounding Madagascar is O. ravaka known from Île Sainte Marie (Dransfield & Beentje 1995).

In both Malesia and Madagascar Orania occupies a great variety of habitats from lowland coastal swampy heath forest about 10 m above sea level to highland tropical humid rainforest about 1220 m above sea level. It also occurs on a wide range of soil types.

So far Orania has never been recorded to have any important economic or commercial value except for its potential ornamental use. Traditionally, however, it has been used by people in many places throughout Malesia and Madagascar for many purposes related to their everyday needs. The trunk is used for building materials in Sumatra, Malay Peninsula and Java (Mogea 1991). Similarly, in Madagascar the wood is also used for house walls and hut construction (Dransfield & Beentje 1995). In New Guinea Orania is more important than in other parts of Malesia and Madagascar. Besides the similar use of wood, the leaves are used for house thatching and the outer wood is commonly used in making arrows and arrowheads (Powell 1976; Zona 1995 pers. comm.).

Although the cabbage and fruit are believed to be poisonous and have never been recorded as normally consumed, their lethality, as mentioned by Burkill (1935) is likely to be just a myth. A person once ate the fruit of Orania regalis planted in Bogor Botanical Garden and had stomach uneasiness and dizziness in the night but soon recovered on the following day. The people of Siwi-Ransiki (Papua) said that the fruit of O. palindan tastes bitter and is frequently consumed by wild pigs without killing either the people or animals. Dransfield & Beentje (1995) reported that Beentje drank some fruit sap of O. longisquama without any ill effects.

Unfortunately, despite its easily recognised appearance, widespread and interesting disjunct distribution (Map 1) and traditionally important uses, Orania has long been one of the least known and most poorly understood members of the Arecoideae.

Fig. 1

Orania tabubilensis. A habit; B diagram of leaf; C mid and apical leaflets; D portion of inflorescence; E triad of flowers on rachilla; F staminate flower, one petal removed; G pistillate flower in section; H fruit; J fruit in section. Scale bar (at C): A 1.6 m; B 1.3 m; C, D 8 cm; E 7 mm; F, G 4 mm; H, I 2 cm. From Baker et al. WJB 1129. drawn by lucy t. smith.

Fig. 2

Orania dafonsoroensis. A mid and apical leaflets; B inflorescence; C staminate flower bud; D staminate flower bud, one petal removed; E pistillate flowers on rachilla; F pistillate flower in longitudinal section; G fruit. Scale bar: A, B 8 cm; C, D 4 mm; E 1 cm; F 5 mm, G 3 cm. From Desianto BD12. drawn by lucy t. smith.

Fig. 3

Orania timikae. A habit; B mid and apical leaflets; C inflorescence; D triad on rachilla; E staminate flower in vertical section; F pistillate flower; G pistillate flower in vertical section. Scale bar: A 1 m; B, C 8 cm; D 1 cm; E, F, G 5 mm. From Dransfield et al JD7667. drawn by lucy t. smith.

Fig. 4

Orania ferruginea. A leaf apex; B inflorescence; C immature fruit; D mature fruit. Scale bar: A, B 9 cm; C 7 mm; D 2.5 cm. From A. P Keim AK 42. drawn by lucy t. smith.

Fig. 5

Orania littoralis. A mid section of leaf; B portion of inflorescence; C staminate flower; D staminate flower, one petal removed; E pistillate flower; F pistillate flower in longitudinal section; G fruit in vertical section. Scale bar: A, B 8 cm; C, D, E, F 4 mm; G 2.5 cm. From Barfod 456. drawn by lucy t. smith.

Fig. 6

Orania subdisticha. A habit; B mid and apical leaflets; C inflorescence; D staminate flowers on rachilla; E staminate flower, one petal removed; F pistillate flowers on rachilla; G pistillate flower in longitudinal section. Scale bar: A 1 m, B, C 8 cm; D 7 mm; E 2.5 mm; F 1 cm; G 5 mm. From Baker et al. WJB 1116. drawn by lucy t. smith.

Fig. 7

Orania bakeri. A habit; B leaflets; C portion of inflorescence; D pistillate flowers on rachilla; E longitudinal section of pistillate flower; F fruit. Scale bar: A 2 m; B 8 cm; C 6 cm; D 2 cm; E 1 cm; F 2.5 cm. A from a photograph, B – F from Baker et al. WJB 581. drawn by lucy t. smith.

Fig. 8

Orania grandiflora. A apex of leaf; B portion of inflorescence; C triads on rachilla, in bud; D pistillate flowers on rachilla; E pistillate flower in vertical section; F staminate flower bud, one petal removed; G double-seeded fruit; H fruit in section. Scale bar A, B 8 cm; C, E 1.5 cm; D 2 cm; F 7 mm; G, H 4 cm. From Heatubun CH179. drawn by lucy t. smith.

Fig. 9

Orania deflexa. A leaf; B mid-leaf leaflets; C inflorescence branch; D pistillate flowers on rachilla; E longitudinal section of pistillate flower; F staminate flowers on rachillae; G staminate flower, one petal removed. Scale bar: A 1 m; B 8 cm; C 6 cm; D, F 1.5 cm; E 7 mm; G 5 mm. From Baker et al. WJB 1319. drawn by lucy t. smith.

Fig. 10

Orania longistaminodia. A apex of leaf; B inflorescence; C staminate flower bud; D staminate flower bud, one petal removed; E pistillate flower; F pistillate flower, one petal removed; G petal. Scale bar: A 6 cm; B 4 cm; C, D 5 mm; E, F, G 7 mm. From Barfod 374. drawn by lucy t. smith.

Fig. 11

Orania zonae. A habit; B portion of leaf sheath; C tip of peduncular bract; D portion of inflorescence; E staminate flower bud; F staminate flower bud, one petal removed; G androecium. Scale bar A 1.5 m; B, C 9 cm; D 6 cm; E, F 4 mm; G 3 mm. From Zona et al. 674. drawn by lucy t. smith.

Map 1

The disjunct distribution of Orania (●).

One difficulty faced by previous workers on Orania was sorting out the nomenclature (i.e. typification) for some of the species. This was presumably due to limited access to type specimens, mainly because some of them have been lost. Furthermore, many collections are incomplete, resulting in doubtful species delimitation. In this study we have had type specimens from various herbaria available. We have also had access to more herbarium specimens than previous workers and have conducted extensive fieldwork and have therefore been able to make more complete morphological observations.

The greatest difficulty in the area of biogeography is the problem of under-collecting. Little collecting of Orania has been done in the eastern part of Indonesia including the Moluccas and Papua. Although there is photographic evidence of Orania in Sulawesi (Beckwith 1940 pers. not. in Fairchild 1943), no collection has ever been made. We have had the advantage of being able to make new collections of Orania in Papua without facing many of the difficulties encountered by previous workers. The results of this fieldwork give new data on the distribution, variation within species in the wild and relationships among the species. This has led to a complete change in the way we look at the distribution of the genus, especially in Malesia.

Finally, all of these factors have provided a basis for an improved working taxonomy and biogeography of the genus.

Morphology

Habit. All species of Orania are solitary, single-stemmed tree palms. Most are massive, with trunks more than 10 cm diam. and tall and sometimes even dominate the canopy. Among these species, O. palindan is the most robust and tallest, with a trunk that can reach 40 cm in diam. and 40 m in height, thus making it very conspicuous in the wild. Eight species, all New Guinean, are slender, with trunks about 10 cm or less in diam. and up to 15 m tall only, thus less conspicuous in the forest or understorey. These species are O. archboldiana, O. ferruginea, O. littoralis, O. oreophila Essig, O. parva Essig, O. subdisticha, O. tabubilensis, and O. timikae, with O. oreophila being the most slender.

Leaves. Leaves of Orania are relatively large, paripinnate and marcescent. All species except O. deflexa have their leaves arching upwards. In O. deflexa the petioles are strongly decurved (arching downwards) giving the impression that the crown is drooping. This phenomenon is unique to the species and thus a very useful field identification character.

Almost all species have their leaves arranged spirally, but four species, Orania disticha (New Guinea), O. ravaka (Madagascar), O. tabubilensis (New Guinea) and O. trispatha (Madagascar) have distichous leaves. Two species have their leaves arranged subdistichously, O. subdisticha and O. timikae. In other species the distichy is inconsistent. Dransfield & Beentje (1995) reported that juveniles of O. longisquama could bear apparently distichous leaves at an early stage. O. dafonsoroensis has subdistichous leaves when juvenile, but these gradually change to spirally arranged leaves in mature specimens. Thus in Orania leaf arrangement may change with maturity.

Leaflets are generally similar throughout Orania. They are single-fold, reduplicate, large, elongate-lanceolate in form, praemorse at the apex and with entire margins. The adaxial surface is usually glabrous and green in colour. The abaxial surface is almost always covered, either sparsely or densely, with a white indumentum or red-brown tomentum. Only in the leaflets of O. tabubilensis is the indumentum sometimes absent.

All species except Orania archboldiana, O. deflexa and O. tabubilensis have their leaflets arranged on the rachis in one plane. In these three species the leaflets are arranged in more than one plane, giving the whole leaf a plumose appearance. However, unlike O. archboldiana and O. tabubilensis, in O. deflexa the irregularity in leaflet arrangement is clearly observed in the middle part of the rachis, while in the distal $\frac{1}{3}$ of the rachis the leaflets are more or less regular in arrangement.

The number of leaflets in the most proximal part of the rachis also varies. Most species have a single leaflet in that location. Four species, Orania grandiflora, O. lauterbachiana, O. longisquama and O. oreophila, have two leaflets in a group, whereas O. archboldiana, O. deflexa and O. gagavu Essig have more than two. It is still unclear whether these two features are significant in the classification of species of Orania but they are important for identification.

Although the adaxial surface of the leaflet is usually glabrous, some species have conspicuous wax, for example Orania decipiens Becc., O. disticha, O. glauca Essig and O. longisquama, with O. glauca being the most glaucous of all.

In addition to the white indumentum or red-brown tomentum, reddish to dark brown ramenta may also be found on the abaxial surface. They are found along veins, mostly on the main vein. In Orania ramenta are restricted to the Madagascar taxa. The petiole and leaf-sheath are similar in all members of Orania. The petiole is conspicuously shorter than the lamina and covered, either sparsely or densely, with tomentum. Although the petiole may be very short, it is still well distinguished from both lamina and leaf-sheath. The leaf-sheath is well-developed, thick, woody and disintegrates into fibres when old, with a glabrous adaxial surface and densely covered with white indumentum and red-brown tomentum on the abaxial surface. In large members of the genus the leaf-sheath is extremely robust. A ligule is present but usually disintegrates into fibres as the leaf becomes old.

Inflorescences. The basic structure of the inflorescence is the same in all species. They are axillary, solitary in the leaf axil, interfoliar, bisexual and protandrous. Flowering is pleonanthic, i.e. it continues throughout the adult life of the palm and does not result in stem death.

In most species of Orania the inflorescence branches are widely spread. In the ‘Sylvicola type’ inflorescence, found in O. sylvicola and O. glauca, there are relatively numerous rachillae, each slender, short and branching at a convergent angle. This type of inflorescence is believed to arise when development of the inflorescence, particularly the rachillae, is restricted by the peduncular bract, which covers the inflorescence during its early development. O. regalis has a congested inflorescence, which may be the most extreme form of a ‘Sylvicola type’ inflorescence.

The number of orders of branching also varies between species. Most species have their inflorescence branching to two orders. Orania dafonsoroensis, O. parva and O. timikae, all from New Guinea, have inflorescences branched to one order only. Five species, three Malesian (O. deflexa, O. gagavu and O. glauca) and two from Madagascar (O. longisquama and O. trispatha) have inflorescences branched to three orders.

The results of this study indicate that variation in the order of branching within species is encountered only in Orania paraguanensis Becc. and O. trispatha. Variation in numbers of orders of branching is not unusual in palms. Dransfield & Beentje (1995) were apparently the first to record its occurrence in Orania. However, as this variation only occurs in the two species, it can still be applied for field identification but is of less importance in classification.

At anthesis the inflorescence is usually 80 to 200 cm long. It can exceed 200 cm in seven species, Orania dafonsoroensis, O. decipiens, O. gagavu, O. glauca, O. macropetala, O. micrantha and O. palindan. Undoubtedly the most robust of all is O. palindan, especially the individuals found in New Guinea, the inflorescences of which can reach lengths of almost 300 cm (Baker 2000 pers. comm.). Only one species, O. longistaminodia, is likely to have a mature inflorescence less than 80 cm long; however, this information is based on a single herbarium specimen (Barfod 374). The total length of the inflorescence of this specimen is approximately 74.5 cm (peduncle about 38.5 cm, rachis about 36 cm). The field notes accompanying the specimen describe the inflorescence as having a peduncle 33 cm long and rachis 20 cm long.

Most species have a robust peduncle more than 1.5 cm diam. except for a group of seven species (Orania archboldiana, O. ferruginea, O. micrantha, O. littoralis, O. oreophila, O. parva and O. timikae), which have slender peduncles 1 – 1.5 cm diam. The peduncle itself is either glabrous or covered, sparsely or densely, with indumentum. Although it is useful in field identification, this character is probably not particularly important as it may vary within species, such as in O. parva and O. palindan. Two species, O. glauca and O. longisquama, have peduncles covered with wax. In O. glauca the wax covers the whole inflorescence, giving a glaucous appearance. This is undoubtedly a useful character in field identification. In most species the peduncle is shorter than the rachis. In O. dafonsoroensis, O. parva and O. timikae the peduncles are two or more times longer than the rachis. This gives the inflorescence a very characteristic appearance.

In Orania, the peduncular bract is the main protective organ for the inflorescence. The prophyll seems to play a much less important role in protecting the developing inflorescence, even though at a very early stage of its development it encloses the entire inflorescence. In all species the peduncular bracts are borne at the base of the peduncle. Malesian species always have a single peduncular bract whereas Madagascar species sometimes have two peduncular bracts. Observations on living specimens of O. regalis in Bogor Botanic Garden during this study showed that the peduncular bract opens by splitting along its length to expose the inflorescence slightly before anthesis and remains in place long after pollination and even until the fruits are mature.

All species of Orania except O. parva have numerous zigzag rachillae in their inflorescence. The rachillae themselves are either slender (diam. less than 0.5 cm) or thick (diam. 0.5 cm or more).

Flower clusters. In Orania, flowers are borne in triads, at least near the base of the rachillae. The one pistillate flower is always located adaxially to the two staminate flowers. Distally the rachillae bear solitary or paired staminate flowers.

Staminate flowers. Most species of Orania have staminate flowers with six or seven stamens, so the number of stamens is not useful in defining most species. The reduction in the number of stamens in O. regalis always takes place on the outer whorl, where the three antesepalous stamens are aborted reducing the number to three.

Amongst the Malesian taxa, Orania macropetala has the highest number of stamens (9 to 14). Another Malesian species, O. zonae, has stamens which vary greatly in number from six to nine or 12 in staminate flowers from the same rachilla. In this species it is very rare to find a staminate flower with six stamens. On close examination the anthers are always free in flowers with six stamens, but united partially or entirely when there are more than six. In flowers with nine stamens, six anthers are united in pairs giving the appearance of six stamens. In flowers with 12 stamens, the anthers are all united in pairs, again giving the appearance of six stamens.

Within the Madagascar taxa Orania longisquama has 16 to 20 stamens while O. ravaka and O. trispatha have more than 20 stamens. In all species except O. regalis and O. zonae the anthers are free. The anthers of O. zonae have already been described. In O. regalis the staminate flowers with three stamens always have free anthers. In flowers with five stamens, two anthers are usually united, thus giving an appearance of three stamens.

In almost all members of Orania a pistillode is usually absent. In O. longistaminodia there is occasionally a prominent pistillode 2 – 2.5 mm long and 3 – 3.5 mm wide. Essig (1980) noted that the staminate flower of O. sylvicola possesses a conspicuous pistillode. Our observations on the herbarium specimens available, including the type specimen (HEJC 1850), do not indicate any pistillode.

Pistillate flowers. Almost all species have more than six pistillate flowers per rachilla. The pistillate flowers are mainly regularly attached to the rachilla with at least 1 cm between two successive pistillate flowers. Only in some individuals of Orania lauterbachiana an irregular pattern of attachment with less than 1 cm distance between adjacent pistillate flowers can be found.

The number of staminodes in almost all species is always six. In Orania regalis three to five are recorded while in O. macropetala up to 10 staminodes can be observed, and in O. longisquama 11 to 20, and in O. ravaka and O. trispatha more than 20 staminodes. Although O. zonae has a variable number of stamens from six, nine and 12, the number of staminodes is always six.

The form of the staminode is uniform in almost all species but in Orania longistaminodia and O. sylvicola at least two staminodes are larger than the rest. The gynoecium is uniformly triloculate and triovulate. All species of Orania have lateral and hemianatropous ovules.

Fruits. Fruits in Orania develop from one, two or three carpels to form unlobed, or two-lobed or three-lobed fruits with sub-basal stigmatic remains. At the base of unlobed fruits there are two small lobes, derived from the two undeveloped or abortive carpels, near the stigmatic remains. In two- or three-lobed fruits, the fruit lobes correspond with the number of functional ovules, in which each lobe is always globose or spherical in most of the species and each of the three carpels has a separate endocarp. Only one species, O. decipiens, possesses sub-pyriform fruits.

Most of the species produce medium to large fruits, 3 – 5.5 cm in diam. Orania regalis has the largest fruit in the genus (about 8 cm in diam. based on the recently collected materials from the Wandammen Peninsula in Papua; Baker 2000 pers. comm.). Fruits are usually dull greenish yellow-brown when mature. Bright orange to reddish orange fruits are found only in the New Guinean species O. dafonsoroensis, O. disticha, O. glauca, O. grandiflora, O. lauterbachiana, O. longistaminodia, O. palindan and O. regalis.

Other features of fruit morphology are generally uniform throughout the genus. Most species have a mesocarp slightly 2 – 4 mm thick. The thickness of the mesocarp was used to separate taxa or even as a basis for new taxa by Beccari (1919a, b). We regard it as less significant as shown in O. decipiens and O. palindan.

The mesocarp may have a significant role in dispersal. The mesocarp in Orania is fibrous but, unlike the mesocarp in Cocos, it is also fleshy, thus may be attractive to wild pigs (Sus sp.), cassowaries (Casuarius sp.) or other animals. Each seed is enclosed in its own hard endocarp. In every location in Papua where we conducted field observations, the indigenous people always mentioned that large mammals and birds are known to consume the fruits. Hornbills and bats were said to eat ripe fruits while still on the tree. Orange or red fruits on green axes exhibit a syndrome associated with dispersal by birds (Van der Pijl 1969). However, it seems unlikely that hornbills will be able to swallow fruit as big as those of Orania. Dispersal by bats seems very unlikely, as bats do not recognise fruits through colour. We have not seen bats eat these fruits, although bats were regularly seen around the infructescences. The large, dull greenish or yellowish fruits could be dispersed by flying foxes, especially Rousettus sp. and Pteropus sp., which are present throughout Malesia (see Musser 1987), but we have no primary sources of information.

In most cases, fallen fruits of Orania are found abundantly and relatively untouched near the parent tree. Rotting fruits of Orania produce a distinctive smell, similar to the smell produced from rotting fruit of mangoes. This can be easily recognised in both day and night by large forest floor animals such as wild pigs (Sus scrofa) and cassowaries (Casuarius sp.) in New Guinea. Wild pigs seem to be common consumers of the fruit; the people in Dungus Iwul in West Java also report that wild pigs occasionally eat fallen fruits of O. sylvicola.

Wild pigs are indigenous in West Malesia, but introduced in New Guinea where they appear to be the feral descendants of pigs carried from island to island by humans in prehistoric time (Maa 1982). The other three species of Artiodactyla present in New Guinea, axis Deer (Axis axis), Java deer (Cervus timorensis) and fallow deer (Dama dama), are more recently introduced, in the late 19th or early 20th Centuries (Ziegler 1982). Therefore, the original agent of dispersal for Orania in New Guinea is probably the cassowary, which is the only large forest floor animal indigenous to the island. Cassowary as agent of dispersal for one species of Orania, O. regalis (then as O. aruensis) was observed by Beccari (1877) on Wokam Island, an island within the Aru Archipelago. As in New Guinea, the cassowary is indigenous to this island.

The other possibility is dispersal by water. Field observations in several areas in Papua, particularly at Nuni-Asai and Siwi-Ransiki, show that individuals of Orania sometimes are found growing near to either rivers or creeks.

Seeds. Seeds in all species of Orania are uniform. The endosperm in Orania is always homogenous with a hollow inside containing liquid endosperm, which still persists in mature fruits.

The surface of the seeds is somewhat grooved by a sparse network of fibres. These fibres are actually anastomosing raphe branches. The raphe itself is a ridge or depression on the seed which usually becomes the source of fibrovascular bundles (Dransfield & Beentje 1996).

A very thin inconspicuous brown testa encloses the seed. Like the endocarp, the testa is also grooved by fibres. All species have a sub-basal embryo except Orania decipiens which has an eccentrically apical embryo.

Seedlings. Germination in all species of Orania is remote tubular. All species possess a bifid eophyll except O. paraguanensis and O. trispatha, which both have a pinnate eophyll.

Taxonomy

A morphological species concept is followed in this monograph, in which delimitation is based on perceived discontinuities in morphological variation (Dransfield 1999). The decision to adopt the morphological species concept is based on the practical nature of the concept. However, the limited number of herbarium specimens affects observations on discontinuities in morphological variations. Under-representation of Orania in herbaria remains a serious problem for the monographer. Within Orania there are two groups of species: those from Madagascar and those from Malesia. These groups do not have any formal taxonomic rank. The species in Madagascar differ from those in Malesia in three characters as can be seen in Table 2. For details on the subgroups and each of the species included, see Taxonomic Account.

Table 2 Differences between species of Orania in Madagascar and those in Malesia.

Taxonomic account

Description of the genus Orania

Orania Zipp. in Blume (1829: 297). Type: O. regalis Zipp.

Arausiaca Blume (1836: 8, t. 119, 122). Type: A. excelsa Blume.

Macrocladus Griff. (Griffith 1845: 489). Type: M. sylvicola Griff.

Sindroa Jum. (Jumelle 1933: 11). Type: S. longisquama Jum.

Halmoorea J. Dransf. & N. W. Uhl (Dransfield & Uhl 1984: 164). Type: H. trispatha J. Dransf. & N. W. Uhl.

Habit solitary, pleonanthic, monoecious palms of the forest understorey and canopy. Stems varying in height, often conspicuously ringed with leaf scars, sometimes with wax. Leaves massive in many species, almost always spirally arranged, rarely distichous or subdistichous, reduplicately pinnate and deciduous under their own weight; crownshaft absent; sheath well developed, thick and woody, splitting longitudinally opposite the petiole, usually densely tomentose, fibrous along the margins; petiole usually short, channelled adaxially, rounded abaxially; rachis much longer than the petiole, curved, abaxially rounded, adaxially angled but channelled near the base; leaf-sheath and ligule usually disintegrating into fibres. Leaflets single-fold, regularly arranged and held in one plane except in three species, O. archboldiana, O. deflexa and O. tabubilensis where the leaflets held in more than one plane giving the whole leaf a plumose appearance, apices praemorse; adaxial surface bright green when mature, glabrous, sometimes with conspicuous wax, abaxial surface densely covered with white indumentum, in Madagascar species usually with ramenta along the main vein, ramenta absent in Malesian species, midrib prominent adaxially; marginal nerves occasionally prominent, transverse veinlets obscure or occasionally conspicuous. Inflorescence axillary, solitary, interfoliar, protandrous, almost all spreading, rarely congested, branched from 1 to 3 orders, most species to 2 orders; peduncle stout, circular in cross-section, covered in caducous brown scales, sometimes greatly elongate to two times or more length of rachis; prophyll persistent, short, tubular, 2-keeled, included within the subtending leaf, usually becoming frayed distally; peduncular bract one; or in Madagascar species sometimes two, borne just above the prophyll, very large and conspicuous, almost woody, tubular, often inflated, completely enclosing the inflorescence in bud, splitting along its length to expose the inflorescence, with a solid, flattened, lanceolate beak, tomentose, eventually deciduous; rachis usually longer than peduncle, brown tomentose and bearing spirally arranged low glabrous, coriaceous, collar-like bracts subtending first-order branches; first-order bearing few, spirally arranged, similar bracts, each subtending 2nd order branch; first-order branches often with a basal pulvinus; further branches subtended by an inconspicuous triangular bract; rachillae spreading, flexuous, bearing rather distant triads proximally and solitary or paired staminate flowers distally; triads superficial, subtended by a minute, scale-like triangular rachilla bract, or rachilla bracts sometimes absent; floral bracteoles minute, scale-like or sometimes absent. Flowers superficially rather similar, cream-coloured, borne in triads in up to half length of rachilla, the rest in dyads, most species with 20 to more than 100 flower clusters per rachilla. Staminate flowers asymmetrical, narrower than the pistillate; sepals 3, almost distinct or more usually united partly; petals 3, fleshy, distinct, valvate, thickened at the apex, lanceolate-elongate or somewhat spatulate; stamens 3 – 14 in Malesian species, 16 – 42 in Madagascar species; filaments dark brown, short to moderate, rather fleshy, distinct or wholly or partially united; anthers creamy pale yellow, lanceolate-elongate, basifixed, erect, with large connective, extrose or latrose, free, occasionally united; pollen elliptic or circular, monosulcate, with fine reticulate or rugulate, tectate or semi-tectate exine; pistillode absent, rarely obvious in O. longistaminodia. Pistillate flowers regularly or slightly irregularly arranged, conical or pyramidal; calyx flattened, short, basally tubular with 3 low, triangular lobes; petals 3, valvate, distinct, triangular; staminodes usually creamy pale yellow, mostly 3 – 6 (–10) in Malesian species, 11 to more than 20 in Madagascar species, very short, awl shaped or well-developed, in most species uniform in size and shape; gynoecium dark brown, triloculate-triovulate, pyramidal; stigmas 3, short, recurved at anthesis; ovule laterally attached, pendulous, presumably hemianatropous. Fruit green in juvenile, greenish yellow or bright reddish orange when mature, developing from 1, 2 or 3 carpels, rounded bi-, or tri-lobed; spherical or globose in one lobed form, rarely obovoid or sub-pyriform; stigmatic remains sub-basal; epicarp smooth, very thin; mesocarp fleshy, traversed by numerous short radial fibres; endocarp dark brown, thinner than mesocarp, with deeply grooved surface, heart-shaped button present basally; testa very thin, dark brown, attached to seed, with grooved surface. Seed spherical, basally attached with a circular hilum, surface somewhat grooved by a sparse network of fibres, endosperm homogenous, sometimes with hollow; clear liquid endosperm present up to maturity, embryo sub-apical or lateral. Germination remote-tubular. Eophyll bifid with praemorse apices, or pinnate. Cytology based on study by Eichhorn 1957, n = 16 (O. sylvicola as O. macrocladus, O. palindan as O. philippinensis); for other species unknown.

Key to the species of Orania

• 1. Ramenta present (Madagascar)...............................................2

  Ramenta absent (Malesia)...........................................................4

• 2. Leaves spirally arranged; pistillate flowers numerous, more than 6 in a rachilla...............................1. O. longisquama

  Leaves distichously arranged; pistillate flowers few, 6 or less in a rachilla........3

• 3. Inflorescence branching to 2 orders; pistillate flowers always less than 6, petals 10 mm long or less; peduncular bract varying from one to two in the same population; eophyll always bifid............2. O. ravaka

  Inflorescence branching to 3 or 4 orders; pistillate flowers up to 6 per rachilla; petal more than 10 mm long; peduncular bracts always two; eophyll sometimes pinnate............................3. O. trispatha

• 4. Inflorescence branching to one order only.......................................5

  Inflorescence branching to more than one order.......................................................7

• 5. Small palm, diam. of trunk around 10 cm or less; number of rachillae in one inflorescence 10 or fewer; bearing triads in the proximal $\frac{1}{5}$ to $\frac{1}{3}$ from the base of rachillae.......................6

  Large palm, diam. of trunk more than 10 cm; number of rachilla in one inflorescence more than 10; bearing triads in the proximal $\frac{4}{5}$ to $\frac{9}{10}$ from the base of rachillae.............................9. O. dafonsoroensis

• 6. Leaves conspicuously spirally arranged, peduncle of inflorescence glabrous, number of flower clusters 120 to 128 per rachilla..................................................................................11. O. parva

  Leaves arranged subdistichously; peduncle of inflorescence with dense red-brown tomentum, number of flower clusters 80 to 85 per rachilla 10. O. timikae

• 7. Inflorescence branching to 3 orders...............................................8

  Inflorescence branching to 2 orders..........................................................10

• 8. Leaves arching upwards; leaflets arranged in one plane........................................9

  ..............Leaves arching downwards (decurving); leaflets arranged in more than one plane......................25. O. deflexa

• 9. Wax absent in inflorescence; rachillae about 30 to 60 cm long; number of flower clusters per rachilla numerous, 80 to 90..........................................................26. O. gagavu

  Wax abundant in inflorescence; rachillae short-branching in convergent angle, about 13 to 15 cm long; number of flower clusters per rachilla few, 20 or less................................................................22. O. glauca

• 10. Leaves in crown not spirally arranged..........................................................11

  Leaves in crown spirally arranged..........................................................13

• 11. Leaves distichously arranged..................................................12

  Leaves sub-distichously arranged..........................................16. O. subdisticha

• 12. Small understorey, trunk c. 5 m high, stem c. 10 cm diam.; leaflets arranged in more than one plane, absent in middle of rachis; inflorescence densely covered with red-brown tomentum..........................................8. O. tabubilensis

  Taller species, trunk 15 – 20 m high, stem 20 – 23 cm diam.; leaflets arranged in one plane, not absent in middle of rachis; inflorescence glabrous........................................................7. O. disticha

• 13. Leaflets arranged in one plane..........................................14

  Leaflets arranged in more than one plane........................................12. O. archboldiana

• 14. Small understorey palm, diam. of trunk always 15 cm or less..................15

  Medium to large palm, diam. of trunk always more than 15 cm..................17

• 15. Number of flower clusters per rachilla 100 or more..................13. O. ferruginea

  Number of flower clusters per rachilla fewer than 100.....................................16

• 16. Length of rachilla 40 cm or longer; species of seacoast and lowland forest, below 1000 m altitude.........................................14. O. littoralis

  Length of rachilla 25 cm or shorter; species of highland/mountain forest, 1000 m or higher altitude......................15. O. oreophila

• 17. Inflorescence congested; number of stamens 3 – 5, never more than five.....................................23. O. regalis

  Inflorescence well expanded; number of stamens more than five..................18

• 18. Number of stamens 6 – 7 or varies 6, 9 or 12 in different flowers but of the same rachilla or inflorescence..................................................................................19

  Number of stamens always 9 – 14.........................................21. O. macropetala

• 19. Filaments always free..................................................................................20

  Filaments always united..................................................................................28. O. zonae

• 20. Staminodes unequal, at least two being different.........................................21

  Staminodes always similar in size and shape..............................................22

• 21. Rachillae branching in convergent angle; staminodes two larger and longer with hooked tips.........................................24. O. sylvicola

  Rachillae not branching in convergent angle; staminodes two or three, larger and longer without hooked tips..................................................................................27. O. longistaminodia

• 22. Species found only in the Philippines..................................................................................23

  Species found in the Philippines, presumably in Sulawesi and East Malesia.........................................24

• 23. Large palm, trunk 18 to 20 m tall, stem 25 – 30 cm diam.; fruit globose; embryo placed below the middle line of the seed; eophyll pinnate..................................................................................6. O. paraguanensis

  Medium to large, trunk up to 8 m tall, stem 10 – 24 cm diam.; fruit sub-pyriform; embryo placed not very far from its summit (eccentrically apical); eophyll bifid.........................................4. O. decipiens

• 24. Length of petal in pistillate flower 10 mm or more.........................................25

  Length of petal in pistillate flower usually less than 10 mm.........................................26

• 25. Rachillae robust, 7 – 8 mm diam., obviously zigzagging, densely covered with red-brown tomentum; mature fruits reddish orange, around 5 cm diam.......................................................................................................20. O. grandiflora

  Rachillae less robust, 5 – 6 mm diam., not obviously zigzagging, with greyish white indumentum; mature fruits dark brown, around 4 cm diam.........................................19. O. bakeri

• 26. Inflorescence glabrous or very rarely with red-brown tomentum; length of petal in staminate flower usually less than 10 mm – very rarely 10 – 11 mm; mature fruit large, 6.5 – 7.5 cm diam..........................................5. O. palindan

  Inflorescence densely covered with red-brown tomentum; length of petal in staminate flower always 10 mm or slightly more; mature fruit 3.5 – 5.0 cm diam.........................................27

• 27. Triads borne in the proximal half of rachilla; staminate flowers blunt c. 10 × 5 mm; lowland species, found at less than 1000 m altitude..................................................................................17. O. lauterbachiana

  Triads borne up to the proximal $\frac{4}{5}$ part of rachilla; staminate flowers elongate with 10 mm long × 2 – 4 mm wide, only rarely 5 mm; highland species, found at more than 1000 m altitude.............................................18. O. micrantha

The group of species in Madagascar. Dransfield & Beentje (1995) recognised three species of Orania in Madagascar: O. longisquama, O. ravaka and O. trispatha (Table 3). Their taxonomy is confirmed in the present study. Dransfield & Beentje (1995) mentioned that the ramenta are red-brown in O. ravaka but pale grey in O. trispatha. However, the results of this study do not indicate any significant differences in the colour of the ramenta.

Table 3 Comparison of morphological characters in the Orania species occurring in Madagascar.

1. Orania longisquama (Jum.) J. Dransf. & N. W. Uhl (Dransfield & Uhl 1984: 164). Type: Madagascar, Port de l’ Anivo, 1932, Perrier 11937 (holotype P).

Sindroa longisquama Jum. (Jumelle 1933: 11). Type: Perrier 11937 (holotype P).

Large palm. Trunk 7 – 20 m tall, c. 13 – 25 cm diam. (dbh), swollen at the base, internodes c. 7 – 10 cm, brown, scars irregular, c. 3 – 6 cm, brighter coloured, wood extremely hard, yellowish brown, bark pale brown, hard. Leaves 9 – 15 in the crown, spirally arranged, c. 3 m long; leaf-sheath c. 26 – 40 cm long, 10.5 – 30 cm wide, massive, margins disintegrating into fibres, fibres straight, c. 7 – 8 cm long, adaxial surface glabrous, abaxial surface with dense red-brown tomentum; petiole c. 1 – 1.2 m long, 2 – 2.5 cm diam., gradually tapering, sometimes margins disintegrating into fibres; rachis c. 1.67 – 2 m long, c. 2.5 cm diam. in the middle part; leaflets elongate-lanceolate, regularly arranged, c. 55 – 65 on each side of rachis, the proximal 2 leaflets in a pair, the middle part with 2 leaflets crowded c. 2 cm distant, otherwise c. 5 cm distant, c. 68.5 – 84 cm long, 2.5 – 5 cm wide, adaxial surface with thin white indumentum and wax, thin red-brown tomentum on midrib, midrib robust, other ribs less robust, abaxial surface with dense white indumentum, red-brown tomentum on the midrib and margins, midrib robust, other ribs thicker than adaxial, ramenta present, red-brown, mainly on midrib and sparsely on other ribs. Inflorescence spreading, branching to 3 orders, robust, c. 83 – 125 cm long, 45 – 110 cm wide; prophyll persistent, c. 28 – 45 cm, 6 – 11 cm wide, hard, disintegrating into fibres when old, adaxial surface glabrous, abaxial surface with dense red-brown tomentum; peduncle massive, c. 25 – 40 cm long, c. 10 cm diam. in the middle, with dense red-brown tomentum; peduncular bract one or two, woody, persistent, proximal peduncular bract inserted 13 cm from base, c. 55 – 57 cm long, 6 cm wide, adaxial surface glabrous, abaxial surface with dense red-brown tomentum, distal peduncular bract inserted 7 cm from base, c. 60 – 95 cm long, 6 – 7.5 cm wide, adaxial surface glabrous, abaxial surface with dense red-brown tomentum, both peduncular bracts splitting in the middle, disintegrating into fibres when old; rachis c. 60 – 69 cm long; first order branches 10 – 20, c. 5 – 10 cm long, rachilla bracts conspicuous, c. 1.2 cm long, 7.5 – 8 mm wide, second order branches c. 20, c. 1.5 – 3 cm long, rachillae robust, c. 24 – 40 cm long, bearing 88 – 143 flower clusters, bearing triads in the proximal $\frac{2}{3}$ part, the basal part of 2nd order branch with c. 1 – 1.5 cm devoid of flowers, triads c. 1.5 – 2 cm distant. Staminate flowers with calyx of 3 united minute sepals; corolla with 3 free petals, c. 5 – 7 mm long, 2 – 3 mm wide; stamens c. 12 – 18, filaments free, dark brown, c. 0.4 – 1.25 mm long, anthers elongate-lanceolate, pale creamy yellow, c. 2 – 5 mm long; pistillodes absent. Pistillate flowers with calyx of 3 united sepals, c. 1 – 2 mm long; corolla with 3 free petals, c. 5 – 7 mm long, 3.5 – 5 mm wide; staminodes 12 – 18, c. 1 – 2.5 mm long; gynoecium dark-brown, c. 3 – 5 mm long, 3 – 4 mm wide; stigmas 3 elongate, brighter coloured. Fruit globose or bilobed, c. 3.5 – 4.5 cm diam., pale green when young, stigmatic remains sub-basal; endosperm white, with a hollow inside, liquid endosperm said to be sweet-tasting (see Beentje & Andriampaniry 4730). Embryo placed below middle line of seed. Eophyll bifid.

distribution. Madagascar, mainly on the eastern part of the island facing the Indian Ocean. Map 2.

Map 2

The distribution of Orania longisquama (●).

specimens examined. madagascar. Port de l’ Anivo, 1932, Perrier 11937 (P holotype, K! photo only); de Brickaville, envirous d’ Anivorano Kely et d’ Andramvolaly Kely, 1951, Cours 4495 (P, K!). Maroantsetra: Masoala Peninsula, West coast, Hiaraka, hill near village, 10 Oct. 1986, J. Dransfield JD 6375 (K!, TAN); Masoala Peninsula, West Coast, Antalavia, valley of river Ampatra, 24 Feb. 1988, J. Dransfield JD 6479 (K!, TAN). Toamasina: Mananara Biosphere Reserve, 6 km E of Antanambe on Biosphere Road, 5 Oct. 1991, Beentje 4455 (K!, TAN); Betampona RNI, 25 Oct. 1991, Beentje 4493 (K!, TAN); Mananara Avaratra, 10 km W of Antanambe, 13 April 1992, Beentje 4616 (K!, TAN); Reserve de la Biosphere de Mananara-Nord, Antanambe Village, 10 km W of Antanambe, 27 July 1994, Brummitt et al. OUEM 9 (K!, TAN); Mahavelona, Analava Forest, 18 Nov. 1999, J. Dransfield JD 7746 (K!); Betampona, 15 March 2003, Britt AB 4 (K!, TAN). Analalava: Analalava, 18 km NNE of Maromandia, 4 July 1992, Beentje 4706 (K!, TAN). Manakara: 39 km NNE of Manakara near main road, 20 May 1992, Beentje 4678 (K!, TAN); Fianarantsoa, 30 km N of Manakara by main road, 10 Sept. 2007, Rakotoarinivo RMJ 362 (K!, TAN). Ifanadiana: 2 Km N of Ambilandrano, 27 July 1992, Beentje 4730 (K!, TAN).

habitat. Lowland to highland tropical rain forest 50 – 550 m above sea level.

conservation status. Least concern (LC) (M. Rakotoarinivo pers. comm.). Orania longisquama occurs throughout the lowland forests of eastern Madagascar almost to its southern tip, with a wide extent of occurrence of 151,841 km² and area of occupancy. It is sometimes locally abundant. Several of its localities are within protected areas. Being thought to be poisonous, the palm is not harvested for palm heart by local people.

etymology. Long scales, regarding the presence of ramenta.

vernacular names. anivona (Betsimisaraka), ovobola fotsy (Toamasina), sindro (Maroantsetra).

uses. Leaves are used for house thatching, trunk for house building. Apex is said to be poisonous.

notes. Prior to this study Dransfield & Beentje (1995) did not include the number and size of pistillate flowers as distinctive characters in their determination key. One of the specimens examined, Beentje 4493, has a structure extending from the leaf-sheath reaching the distal part of the petiole. In some way it is similar to a ligule, but Orania is otherwise not known to have a ligule. Dransfield & Beentje (1995) explained this as the leaf-sheath being split to give a tongue-like structure, reaching up to the proximal leaflets producing an apparent petiole.

2. Orania ravaka Beentje (in Dransfield & Beentje 1995: 119). Type: Madagascar, Maroantsetra, Andranofotsy river, Sahavary, hill E of Andilampananina village, 23 Oct. 1986, J. Dransfield JD 6401 (holotype K!).

Large palm. Trunk c. 10 – 20 m tall, grossly swollen at base, c. 10 – 20 cm diam. (dbh), internodes 2.5 – 5 cm, greyish brown, obscurely ringed with scars, bark pale brown. Leaves 6 – 8 in the crown — some old individuals with 9 – 13, distichously arranged, densely covered with red-brown tomentum, c. 2.5 – 4.5 m long; leaf-sheath about 21.5 – 35 cm long, 8.5 – 11 cm wide, margins disintegrating into fibres, in some individuals rarely, adaxial surface with thin red-brown tomentum, wax present, abaxial surface with dense red-brown tomentum and creamy white indumentum, sometimes with greyish white indumentum; petiole c. 30 – 94 cm long, c. 1.7 – 2 cm diam. in the middle, margins disintegrating into fibres; rachis about 198 – 321 cm long, c. 2 cm diam. in the middle; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, c. 33 – 44 on each side of rachis, leaflets c. 5.5 – 7 cm distant, in the proximal c. 3 cm, c. 50 – 74 cm long, 4 – 5 cm wide, adaxial surface glabrous with white substance scattered, thin red-brown tomentum on basal part and midrib, greyish white indumentum on midrib, midrib robust, other ribs thick, abaxial surface with dense white indumentum and thin red-brown tomentum, midrib robust, other ribs thick; ramenta present, mainly on midrib, sparsely on other ribs, red-brown or dark brown, conspicuous, c. 1 cm long. Inflorescence spreading, c. 100 cm long, branching to 2 orders, robust; prophyll hard, persistent, margins adaxially disintegrating into fibres, c. 26.5 – 32 cm long, 3.5 – 8 cm wide, adaxial surface glabrous, abaxial surface deeply covered with red-brown tomentum; peduncle curved, c. 60 cm long, with dense red-brown tomentum, wax present; peduncular bract one or two, woody, c. 45 – 62 cm long, 9 cm wide, adaxial surface glabrous, abaxial surface deeply covered with red-brown tomentum; rachis c. 40 cm; first order branches 13, c. 4 – 9 cm long each, rachilla bracts minute, in some individuals conspicuous, c. 1.5 cm long, 0.8 cm wide; rachillae robust, c. 26 – 40 cm long each, bearing 54 – 78 flower clusters, triads arranged in the proximal $\frac{1}{5}$ part, the basal c. 2.5 – 3.5 cm devoid of flowers, triads c. 1.5 – 3 cm distant, rachilla glabrous, number of triads in rachilla 1 – 3. Staminate flowers with calyx of 3 united minute sepals; corolla creamy white, with 3 free petals, c. 9 – 15 mm long, 3 – 5 mm wide; stamens 16 – 23, filaments free, dark brown, c. 1.25 – 1.5 mm long, anthers elongate-lanceolate, c. 5 – 6 mm long, free, pistillode unknown. Pistillate flowers with calyx of 3 united sepals, 4 mm long; corolla creamy white, with 3 free petals, 10 × 8 mm; staminodes c. 20, c. 3 mm long, uniform; gynoecium c. 8.5 – 9 mm long, 4 mm wide, pale brown, stigma with 3 elongate lobes. Fruit globose or bilobed, c. 4 – 4.5 cm diam., in an individual said to have c. 6 – 8 × 4 cm (Du Puy 152), dull olive green when young, yellowish green when mature, stigmatic remains sub-basal; epicarp smooth, thin, c. 0.7 – 0.8 mm; mesocarp fibrous, c. 2 mm thick; endocarp hard, thinner than epicarp; testa light brown, thinner than endocarp, attached to endosperm; endosperm, creamy white, with c. 3 cm diam., with a hollow inside, c. 8 mm diam. Embryo placed below middle line of seed. Eophyll bifid.

distribution. Madagascar, mostly in the eastern part of the island facing the Indian Ocean and on the north coast. Dransfield & Beentje (1995) reported the presence of the species on Île Sainte Marie, an island just off the northeast coast of mainland Madagascar, but no specimen has been collected so far. Map 3.

Map 3

The distribution of Orania ravaka (●) and O. trispatha (◊).

specimens examined. madagascar. Maroantsetra: Masoala Peninsula, west coast, Hiaraka, 11 Oct. 1986, J. Dransfield JD 6376 (K!, TAN); Andranofotsy river, Sahavary, hill E of Andilampananina village, 23 Oct. 1986, J. Dransfield JD 6401 (K! holotype, TAN); Andranofotsy, Sahavary, hill E of village, 11 Feb. 1988, J. Dransfield JD 6458 (K!, TAN); Masoala Peninsula, c. 6 km of village Hiaraka, 21 April 1989, Du Puy 152 (K!, P, TAN); Sahavary, April 1989, Du Puy & Du Puy MB 161 (K!, TAN). Toamasina: Mananara Biosphere Reserve, 10 – 12 km W of Antanambe, 8 Oct. 1991, H. Beentje 4466 (K!, MO, P, TAN); Soanierana-Ivongo, Ambatovaky Resv., 11 Nov. 1999, J. Dransfield JD 7731 (K!, MO, P, TAN); Maroansetra, Makira, Andranomenahely, 14 May 2007, Rakotoarinivo et al. RMJ 338 (K!, MO, P, TAN).

habitat. Varying from lowland up to rather highland tropical rainforest from about 80 up to 550 m above sea level.

conservation status. Vulnerable (VU D1, D2) (M. Rakotoarinivo pers. comm.). Orania ravaka has a restricted distribution in Madagascar and an estimated population size of fewer than 800 individuals with an extent of occurrence of 8913 km² and area of occupancy 220 km². Some of its localities are within protected areas.

etymology. Jewel in Malagasy language of Betsimisaraka dialect.

vernacular names. menavozona (Hiaraka), sindro (Sahavary), vapaka fotsy (Betsimisaraka).

uses. Leaves are used for house thatching and trunk for housing. No information about the use of fruits.

notes. Orania ravaka is the most recently recognised species from the genus Orania in Madagascar (see Dransfield & Beentje 1995).

3. Orania trispatha (J. Dransf. & N. W. Uhl) Beentje & J. Dransf. (Beentje & Dransfield 1995: 118). Type: Moore 9921 (holotype BH).

Halmoorea trispatha J. Dransf. & N. W. Uhl, (Dransfield & Uhl 1984: 166). Type: Moore 9921 (holotype BH).

Large palm. Trunk c. 15 – 22 m tall, c. 20 – 30 cm diam. (dbh), much swollen at base, trunk very hard, bark light brown, internodes 12 cm, scars 5 cm, brighter coloured. Leaves 10 – 12 in the crown, distichously arranged, c. 3.01 – 5 m long, in some individuals also with appearance of brown coloured resin in every section; leaf-sheath massive, c. 20.5 – 60 cm long, 19.5 – 20 cm wide, margins disintegrating into fibres, adaxial surface with thin red-brown tomentum and wax, abaxial surface with dense red-brown tomentum and wax; petiole c. 75 – 200 cm long, c. 2 – 2.5 cm diam., with dense red-brown tomentum, wax present, margins disintegrating into fibres; rachis about 205 – 240 cm long, c. 2.5 – 3 cm diam., with dense red-brown tomentum and wax; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, 60 – 65 on each side of rachis, leaflets c. 4 – 5 cm distant, c. 98 – 110 cm long, 4.5 – 5.5 cm wide, adaxial surface glabrous, with shiny surface, with red-brown tomentum and white indumentum on midrib, midrib robust, other ribs slender; abaxial surface densely covered with white indumentum and thin red-brown tomentum, mainly on basal part and every rib, midrib thick, other ribs slender; ramenta present, mainly on midrib and sparsely on nearby ribs, c. 1 cm long, greyish white. Inflorescence spreading, branching to 3 – 4 orders, robust, c. 1.46 – 1.5 m long, in some individuals with strong fragrance; prophyll persistent, c. 28.5 – 40 cm long, 7 – 12 cm wide, adaxial surface glabrous, abaxial surface with red-brown tomentum, adaxially disintegrating into mass of fibres when old, curly; peduncle c. 16 – 25 cm long, with dense red-brown tomentum; peduncular bracts two, woody, persistent, splitting in the middle, margins disintegrating into fibres, proximal peduncular bract c. 45 cm long, proximally attached c. 18 – 19 cm, adaxial surface glabrous, abaxial surface with dense red-brown tomentum, distal peduncular bract c. 90 cm long, proximally attached c. 7 – 8 cm, adaxial surface glabrous, abaxial surface with red-brown tomentum and white indumentum; rachis 30 – 40 cm long, with dense red-brown tomentum; c. 8 – 10 first order branches, c. 9 – 20 cm long, 2nd order branches c. 6.5 – 9 cm long; 3rd order branches c. 5.5 cm long, rachis bract conspicuous, c. 3.5 × 9 cm; rachillae robust, c. 28 – 35 cm long each, bearing 36 – 42 flower clusters, bearing triads in the proximal $\frac{1}{{10}}$ part, maximum number of triads 6, rarely seen with 6 triads in rachilla, the basal c. 3 cm devoid of flowers. Staminate flowers with calyx of 3 minute united sepals; corolla with 3 free petals, c. 12 × 5 mm wide; stamens 40 – 42, filaments dark-brown, free, c. 1.25 – 1.5 mm long, anthers elongate-lanceolate, pale creamy yellow, 5 – 6 mm long, free; pistillodes absent. Pistillate flowers with calyx of 3 united sepals, c. 2 mm long; corolla with 3 free petals, c. 3 cm long, 1.1 – 1.2 cm wide; staminodes more than 20, c. 7 mm long, uniform; gynoecium c. 20 mm long, 9 – 10 mm wide, stigma with 3 elongate lobes, c. 6 mm long, darker coloured. Fruits globose, bilobed or trilobed, green to yellowish green when mature, c. 4 – 5 cm diam.; epicarp smooth, thin; mesocarp fibrous, c. 3 mm thick; endocarp thinner than mesocarp, hard, red-brown; testa conspicuously thin, attached to endosperm; endosperm white, c. 3 – 3.5 cm diam., with a hollow inside, liquid endosperm present, said to taste bitter (Dransfield & Beentje 1995). Embryo placed below middle line of seed. Eophyll bifid or pinnate (observed in Beentje 4522).

distribution. Madagascar, mainly on the eastern part of the island facing the Indian Ocean. Map 3.

specimens examined. madagascar. Maroantsetra: Andranofotsy river, Sahavary, 23 Oct. 1986, J. Dransfield JD 6400 (K!, TAN). Toamasina: Mananara Reserve de la Biosphere-Nord, Antanambe Village, 10 km W of Antanambe, 28 July 1994, Brummitt et al. OUEM 12 (K!, TAN); Betampona, 15 March 2003, Britt AB 16 (K!, TAN); Soanierana-Ivongo, Manompana, Befanjana, 16 Sept. 2006, Rakotoarinivo et al. RMJ 266 (K!, P, TAN). Farafangana: Manombo, Reserve Speciale, Parc. 1, 13 – 14 Nov. 1991, Beentje 4522 (K!, TAN).

habitat. Varying from lowland up to highland tropical rain forest, commonly found near to rivers or even also found growing in swamp forest, with altitude range from 50 – 400 m above sea level.

conservation status. Vulnerable (VU D1, D2) (M. Rakotoarinivo pers. comm.). Orania trispatha occurs in lowland forest in the area of the Bay of Antongil, with an extent of occurrence of 25198 km² and area of occupancy 1644 km² and a population size estimated at 600 individuals. It is rarely abundant. Some of its localities are within protected areas.

etymology. With three spathes, referring to the prophyll and two peduncular bracts.

vernacular names. anivo (Farafanga), sindro (Maroantsetra).

uses. Leaves are used for house thatching, trunks for building houses and bridges.

notes. One herbarium specimen observed (Beentje 4522) shows a pinnate eophyll, but some others show bifid eophylls.

The group of species in Malesia. There are at least seven sub-groupings of species within the group of Malesian species, two of which we regard as species complexes (Table 4).

Table 4 Sub-groupings within the Malesian species of Orania.

Species in the Philippines

Table 5 provides a comparison between the three members of this subgroup. The difficulty of species delimitation in the Philippines was recognised by Beccari (1919b). Individuals belonging to the same species seldom exhibit constant characters when growing on separate islands, or even in distant localities on the same island. Further studies on the species that occur in the Philippines backed with more extensive fieldwork are needed.

Table 5 Morphological features distinguishing Orania decipiens, O. palindan, and O. paraguanensis.

4. Orania decipiens Becc. (Beccari 1909: 614). Type: Philippines, Mindoro, Bongabong river, April – May 1906, Merritt 4120 (holotype FI!; isotype K!).

Orania decipiens Becc. var. mindanaoensis Becc. (Beccari 1919b: 333). Type: Whitford & Hutchinson 9179 (holotype FI!), synon. nov.

Orania rubiginosa Becc. (Beccari 1919b: 333) (syntypes Curran 17192, Curran 17259, Loher 7090, McGregor 10575, Ramos 13390), synon. nov.

Medium to large palm. Trunk up to 8 m high, c. 10 – 24 cm diam. (dbh). Leaves robust, spirally arranged, c. 3 – 5 m long; leaf-sheath c. 23 – 35 cm long, 4.7 – 8 cm wide in near the base, adaxial surface glabrous, abaxial surface with red-brown tomentum, margins disintegrating into fibres; petiole c. 51 – 73 cm long, c. 4 – 5 cm diam., glabrous, or with dense red-brown tomentum, wax present; rachis c. 126 – 392 cm long, c. 5 cm diam. in middle part, glabrous or with dense red-brown tomentum, wax present; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, in old leaflets margins disintegrating, the proximal part with singular smallest leaflet, c. 34 in each side of rachis, distance between 2 leaflets c. 5 – 8 cm in middle part, c. 1.05 – 1.5 m long, 6 – 7 cm wide, adaxial surface glabrous, with sparsely wax covering, midrib robust, other ribs less robust, abaxial surface densely covered with white indumentum, sparse red-brown tomentum present on margins, wax present, midrib less robust than adaxial, other ribs more slender. Inflorescence spreading, robust, branching to 2 orders, c. 2 – 2.03 m long; peduncle c. 84 – 84.5 cm long, glabrous, or with dense red-brown tomentum, margins in old inflorescence sometimes found disintegrating to form a mass of fibres; peduncular bract one, woody, persistent, abaxially with dense red-brown tomentum, c. 51 – 52 cm long, 5 cm wide, splitting in the middle, disintegrating into fibres when old; rachis c. 1.16 – 1.18 m long; first order branch 40 – 70 cm, rachillae bract minute, rachillae slender, zigzagging distally, c. 32 – 45 cm long, bearing 77 – 105 flower clusters, bearing triads $\frac{1}{3}$ – $\frac{1}{2}$ part of rachillae, the basal c. 1 – 2.5 cm devoid of flowers, triads c. 1 – 1.7 cm distant, obvious pulvinate form in the base of rachilla, rachilla glabrous. Staminate flowers with calyx of 3 united minute sepals; corolla with 3 free petals, c. 6 – 13 mm long, c. 1 – 2 mm wide; stamens 6, filaments dark-brown, free, c. 0.6 – 1.5 mm long, anthers elongate-lanceolate, free, pale creamy yellow, c. 3 – 6 mm long; pistillode absent. Pistillate flowers with calyx of 3 united sepals, c. 3.5 mm long, 2 mm wide; corolla with 3 free petals, c. 6 mm long, 2 – 2.5 mm wide; staminodes 6, c. 0.6 – 0.7 mm long; gynoecium dark-brown, c. 5.5 × 1.5 mm; stigma with 3 slightly elongate lobes, 1.5 mm elongate. Fruit sub-pyriform, c. 3.2 – 4.2 cm diam., stigmatic remains sub-basal, dull green when young, yellowish green when mature; epicarp c. 0.5 mm thick; mesocarp fibrous, c. 2 – 4 mm thick; endocarp hard, red-brown, c. 0.7 – 1 mm thick; endosperm homogenous, white or creamy white, c. 2.2 cm diam., with hollow inside. Embryo placed not very far from its summit (eccentrically apical). Eophyll bifid.

distribution. Widespread species in the Philippines Archipelago, from Luzon Island in the north to Mindanao in the south, excluding Palawan Island. Map 4.

Map 4

The distribution of Orania paraguanensis (▲) and O. decipiens (●).

specimens examined. philippines. Mindanao: Port Banga, Zamboanga, Jan. 1908, Whitford & Hutchinson 9179 (FI!). Basilan: Zamboanga, 5 – 18 Jan. 1941, Ebalo 975 (K!). Mindoro: Bongabong river, April – May 1906, Merritt 4120 (FI! holotype; K!). Luzon: Bicol Nat. Park, Camarines Norte, 25 May 1885, E. S. Fernando 507 (K!); Wakar, Sierra Madre Mts near Sitio Sablang, 1 June 1885, E. S. Fernando 514 (K!); Nakar, Sablang, Sierra Madre Mts, 2 June 1887, E. S. Fernando 706 (K!); Rizal, May 1906, A. Loher 7066 (FI!, K!); Angilog, Rizal, May 1906, A. Loher 7090 (FI!, K!); Cagayan, Feb. 1909, H. M. Curran 17068 (FI!); Cagayan, March 1909, H. M. Curran 17192 (FI!); Cagayan, March 1909, H. M. Curran 17259 (FI!); Cagayan, East Coast, Aug. 1909, McGregor 10575 (FI!); Bicoyan bay, Isabella, Aug. 1909, McGregor 10658 (FI!); Tagcauayan, Tayabas, March 1911, M. Ramos 13390 (FI!); Zambales, Mt Marayep, Dec. 1924, Ramos & Edaño FB 44779 (BO!, K!).

habitat. Commonly found in lowland tropical rainforest. Altitude from about 100 – 800 m above sea level.

conservation status. Vulnerable (VU D2). Although Orania decipiens has a wide extent of occurrence In Luzon and Mindanao, only seven unique localities have been recorded and much of the lowland forest where it occurred has already been destroyed.

etymology. Deceptive, resembling another species.

vernacular names. angilog (Rizal), bakal (Basilan), banga (Dumagat).

uses. Trunk is used for housing; leaves are used for thatch.

notes. Within Orania decipiens, Beccari (1919b) proposed a new variety montana based on differences in size and shape of the fruit and the thickness of the mesocarp. This variety is treated in this study as belonging within O. palindan. In the protologue of var. montana Beccari mentioned it has a height of 35 feet (approximately 10 m) with diam. of trunk 30.5 cm. Although its height is not within the range of O. palindan, it is taller than O. decipiens (Table 5). On the contrary, the diam. is within the range of O. palindan. Beccari also mentioned that the fruit of O. decipiens var. montana is exactly spherical (i.e. globose) with 42 mm in diam., thus resembling O. palindan. The rest of the description of var. montana (Beccari 1919a) matches well with O. palindan. Furthermore, examination of the syntypes of O. decipiens var. montana, including the specimens used by Beccari himself, show that this variety has a completely globose fruit and thus cannot be included in O. decipiens. Thus, var. montana is transferred to O. palindan. Beccari (1919b) also proposed variety mindanaoensis, distinct from other specimens of O. decipiens in two characters (Table 6). Unfortunately, no specimens of var. mindanaoensis have been collected since the type. Nevertheless, the fruit of O. decipiens var. mindanaoensis is smaller than O. decipiens. Observations on cross-sections of two fruits of the holotype showed that the mesocarp was indeed only 2 – 2.5 mm thick, as opposed to 3 – 4 mm thick in O. decipiens. However, we consider these differences very slight and not important. Although we could not observe the embryo position, the type specimen of var. mindanaoensis possesses the sub-pyriform fruit, which characterises O. decipiens (Table 5). Until we have more collections of O. decipiens var. mindanaoensis, this taxon is subsumed. The two obvious distinctive characters possessed by O. decipiens, the sub-pyriform fruit and embryo placed not very far from its summit or eccentrically apical (Table 5), are also found in O. rubiginosa. Beccari (1919b) separated O. rubiginosa from O. decipiens because a dense red-brown tomentum covers the inflorescence in O. rubiginosa whereas the inflorescence in O. decipiens is glabrous. The results of this study (Table 7) show that the holotype of O. decipiens does have a slight red-brown tomentum in its inflorescence and red-brown tomentum varies in abundance and thickness among the specimens examined. Furthermore, the size (represented by the diam.) of O. rubiginosa fruits — missed by Beccari — is not conspicuously different from O. decipiens. We therefore conclude that the density of red-brown tomentum in the inflorescence and the size (i.e. diam.) of fruit do not reliably distinguish O. rubiginosa from O. decipiens. However, Fernando (2001 pers. comm.), on the basis of his field observations, considers that O. rubiginosa is quite distinct from O. decipiens as it is much smaller (stems no longer than 3 m in already fruiting individuals). We regard height as being less important than the shape of the fruits, in which both of the taxa (O. rubiginosa and O. decipiens) have sub-pyriform fruits. Thus O. rubiginosa is subsumed. O. decipiens shares the same habitat with O. palindan and it has the centre of distribution in Luzon Island as well. O. decipiens occurs in Mindanao Island scattered only to the western part of the island and on the small adjacent island, Basilan. This is in contrast with O. palindan for this species is more commonly found in the eastern part of Mindanao Island. Only on Mindoro does O. decipiens occur without co-existence with O. palindan.

Table 6 Morphological differences between Orania decipiens and O. decipiens var. mindanaoensis according to Beccari (1919b).

Table 7 Variation in the density of red-brown tomentum in herbarium specimens of Orania rubiginosa and O. decipiens.

5. Orania palindan (Blanco) Merr. (Merrill 1905: 88). Type: Caryota palindan Blanco, Philippines, Luzon, Montalban, Manila, March 1912, Merrill Species Blancoanae 144 (neotype K!).

Caryota palindan Blanco (1845: 513).

Orania regalis Naves (1883: 279) non Blume. Type: C. palindan Blanco.

Orania philippinensis Scheff. ex Becc. (Beccari 1885: 156, t.14). Type: Giardino Botanico di Buitenzorg, May 1878, Beccari s.n. (holotype FI).

Orania moluccana Becc. (Beccari 1885: 163). Type: Giardino Botanico di Buitenzorg, May 1878, Beccari s.n. (holotype FI), synon. nov.

Orania regalis (non Zipp.) Miq. (Miquel 1868: 4). Type: O. moluccana Becc.

Orania palindan (Blanco) Merr. var. sibuyanensis (Becc.) Merr. (Merrill 1905: 88). Type: Elmer 12066 (holotype FI), synon. nov.

Orania decipiens Becc. var. montana Becc. (Beccari 1919a: 3018) (syntypes: Elmer 13970, Elmer 1181), synon. nov.

Orania philippinensis Scheff. ex Becc. var. sibuyanensis Becc. (Beccari 1919b: 332). Type: O. palindan (Blanco) Merr. var. sibuyanensis (Becc.) Merr.

Large palm. Trunk 20 – 30 m high, c. 15 – 40 cm diam. (dbh). Leaves 10 – 15 in the crown, spirally arranged, massive, c. 2.5 to 3.5 m long, in the old crown c. 5.6 – 6 m long; leaf-sheath c. 28 × 9.5 cm, adaxial surface glabrous, abaxial surface with red-brown tomentum, margins disintegrating into fibres, fibres short, c. 5 cm long; petiole c. 1.5 m long, with red-brown tomentum; rachis c. 2 – 2.5 cm diam., with dense red-brown tomentum; leaflets c. 100 – 122, elongate-lanceolate, regularly arranged, held in one plane, c. 4.5 – 8 cm distant in the middle part of rachis, c. 1 – 2 m long, 4.5 – 7.5 cm wide, adaxial surface glabrous, with thin red-brown tomentum on midrib, midrib robust, other ribs less robust, glabrous, abaxial surface with dense white indumentum, red-brown tomentum on margin and midrib, midrib robust, other ribs less robust, glabrous. Inflorescence spreading, robust, branching to 2 orders, c. 1.15 – 2 m long; prophyll persistent, c. 27 – 68.5 cm long, 7 – 8.7 cm wide near the base, disintegrating into fibres; peduncle 80 – 150 cm long, glabrous or with thin red-brown tomentum; peduncular bract one, woody, c. 53 – 160 cm long, 14 cm wide, adaxial surface glabrous, abaxial surface with dense red-brown tomentum, splitting in the middle, disintegrating into fibres, wax present; first order branches reaching c. 100 – 105 cm long, bracts subtending rachillae c. 5 mm long; rachillae thick, straight or not conspicuously zigzag, c. 30 – 66.5 cm long, bearing 60 – 130 flower clusters, triads arranged in the proximal half part, the basal c. 3.5 – 6 cm devoid of flowers, triads c. 2 – 3.5 cm distant, rachilla glabrous. Staminate flowers with calyx of 3 united sepals, c. 1 mm long; corolla with 3 free petals, c. 4 – 11 mm long, 2 – 3 mm wide; stamens 6, filaments free, dark-brown, c. 0.6 – 0.75 mm long, anthers elongate-lanceolate, pale creamy yellow, always free, c. 3 mm long; pistillode absent. Pistillate flowers with calyx of 3 united sepals, 1.5 mm long; corolla with 3 free petals, c. 3 – 6 mm long, 2 – 4 mm wide; staminodes 6, c. 1 – 1.75 mm long; gynoecium dark-brown, c. 3 – 4 mm long, 3 – 3.5 mm wide, stigma of 3 elongate lobes, 1 mm long. Fruit globose or bilobed, c. 6.5 – 7.5 cm diam., sometimes found 4 to 5.5 cm diam., stigmatic remains sub-basal, olive green when young, yellowish green to bright yellow when mature; epicarp smooth, thin, c. 0.5 mm thick; mesocarp fibrous, 3 – 4 mm thick; endocarp hard, red-brown, thin, c. 0.5 mm thick; endosperm white or creamy white, c. 3.2 – 3.5 cm diam., with a hollow, 1 cm diam. Embryo placed below middle line of seed. Eophyll bifid.

distribution. A widespread species known from Luzon in the northern part of the Philippines, Sulawesi, Maluku, Misoöl Island and northern part of mainland New Guinea. In Papua it is only known from Manokwari, through the Wandammen Peninsula, to an area near Sentani Lake in Jayapura excluding the Cyclops Mountain. In Papua New Guinea, the presence of this species is so far recorded from the Western Province, close to the border with Papua. So far this species has never been recorded in Borneo. Its occurrence in Sulawesi is based only on an entry in a diary with a photograph written and taken by Ned Beckwith in the 1940s (see Fairchild 1943). Map 5.

Map 5

The distribution of Orania palindan (●).

specimens examined. philippines. Agusan: Cabadbaran, Mt Urdaneta, Oct. 1912, A. D. E. Elmer 13970 (BO!, FI!, L!, NY!). Luzon: Zambales, Nov. – Dec. 1915, Curran et al. 8413 (FI!); Tayabas, May 1908, Curran 10410 (FI!); Cagayan, Feb. 1909, Curran 17068 (FI!). Central Luzon: Caypilayan, Lanati-Morong, 25 March 1893, A. Loher 1355 (K!); Montalban, Manila, 5 May 1893, A. Loher 1401 (K!); A. Loher 1402 (K!); 1906, A. Loher s.n. (K!); no date, A. Loher 1404 (K!); no date, A. Loher 5225 (K!); March 1912, Merrill Sp. Blancoanae 144 (BO!, K!, L! neotype); Batangas, Aug. 1914, M. Ramos 1900 (BO!, L!); Rizal, Dec. 1915, M. Ramos 24107 (BO!, FI!, K!, L!). Laguna: Calauan, Nov. – Dec. 1910, R. C. McGregor 12391 (FI!, K!, L!); Mt Maquiling, Los Baños, 6 July 1917, A. D. E. Elmer 18242 (BO!); Mt Maquiling, Los Baños, June – July 1917, A. D. E. Elmer 18248 (K!); Mt Maleiling, Los Baños, 30 Sept. 1985, E. S. Fernando 553 (K!). Benguet: Baguio, March 1907, A. D. E. Elmer 8950 (BO!, K!, L!). Sibuyan: Magallanes, Mt Giting-giting, April 1910, A. D. E. Elmer 12066 (BO!, FI!, K!, L!, NY!); Magdiwang, Mt Giting-giting in Northern slope, 21 May 1987, E. S. Fernando 696 (K!). Romblon: Magdiwang, Bario Hawasan, 26 May 1992, B. C. Stone et al. PPI 6708 (K!). Davao: Todaya, Mt Apo, May 1909, A. D. E. Elmer 11881 (BO!, K!, L!); Davao, March – April 1927, M. Ramos & G. Edaño 49293 (BO!); Camarines Sur, Oct. 1928, G. Edaño 75896 (BO!). INDONESIA. Moluccas. Halmahera: S. O. A. Tobaru, North Halmahera, 5 Aug. 1922, Beguin 2105 (BO!); 23 Aug. 1922, Beguin 2043 (BO!); Ekor Kali Dowora Ina, 24 Sept. 1974, E. F. de Vogel 3082 (BO!, K!, L!); 25 Sept. 1974, E. F. de Vogel 3102 (BO!, K!, L!); Central part, Akelamo Oba, 4 Dec. 1974, E. F. de Vogel 4402 (BO!, K!, L!); South Halmahera, Payahe, 9 April 1983, J. P. Mogea 4469 (BO!, K!, L!); 20 km SE of Dodinga, Tapayo, 13 Sept. 1985, T. C. Whitmore et al. 3674 (K!). Bacan: 1859 – 1860, Teijsmann et al. s.n. (L!); NE of the Island, 2 Sept. 1985, T. C. Whitmore et al. 3576 (K!); Seyoang R., Mt Sibela, N. Bacan, 31 Aug. 1986, Ramlanto 911 (BO!); Jokir (Yokir?), no date, Anonymous s.n. (FI!). Papua Barat. Sorong: Waigama, Misoöl Island, Aug.?, Anonymous s.n. (BO!). Manokwari: Nuni-Asai, 19 March 1999, A. Keim AK 38 (BO!, K!); A. Keim AK 39 (K!); Nuni-Asai, 20 March 1999, A. Keim AK 40 (K!, BO!, MAN!, BH!, NY!, L!); Waren, Siwi-Ransiki, 2 March 1940, Kanehira & Hatusima 12127 (BO!); Near Ransiki, July 1948, Kostermans P (BO!); Siwi-Ransiki, 25 March 1999, A. Keim AK 43 (BH!, BO!, K!, L!, MAN!, NY!); A. Keim AK 44 (BO!, K!, MAN!). Wandammen: Wandammen Peninsula, Wasior Distr., Kowi, near Wondiwoi village, c. 9 km S of Wasior, 23 Feb. 2000, W. J. Baker 1049 (BO, K!, L, MAN). Bintuni: Bintuni, near Tanah Merah village, 14 Feb. 2002, R. A. Maturbongs RAM 716 (BO!, K!, MAN); 15 Feb. 2002, R. A. Maturbongs RAM 724 (AAU, BO!, K!, LAE). Papua. Yapen: Saubeba-Konti Road, E of Serui, 11 Oct. 2006, A. Keim 805 (BO!). Jayapura: Kampung Puay, Sentani, 22 Feb. 1999, A. Keim AK 37 (BH!, BO!, K!, L!, MAN!, NY!); Arso, Tami River, Isobo, Gusbager PG07 (AAU, K!, LAE, MAN). PAPUA NEW GUINEA. Western: N of Fly Distr., junction of Harvey Creek & Ok Mani R., 10 km WNW of Tabubil, 51°4'28.5''S 141°8'22.1''E, 14 Dec. 2000, W. J. Baker 1140 (AAU, BO!, K!, LAE). CULTIVATED. Java: Kebun Raya Bogor, 16 Jan. 1999, A. Keim et al. AK 34 (BO!, K!); 16 June 1999, A. Keim et al. AK 45 (BO!, K!); 16 June 1999, A. Keim et al. AK 46 (BO!, K!). Singapore: Singapore Botanic Garden, 10 Jan. 1933, Kiah 26185 (K!); 22 June 1929, Nur s.n. (K!).

habitat. Lowland to rather hilly tropical rainforest with altitude from about 10 up to 800 m above sea level.

conservation status. Least concern (LC). On a global scale this species is widespread, occurring from Philippines through to New Guinea, a very large extent of occupancy. It is, however, locally threatened in the Philippines.

etymology. Native name in Tagalog Language.

vernacular names. ajabu (Sumuri language), banga (Tagalog-Bagobo and Manobo dialects), barongong (Tagalog-Montalban dialect), hiyaub (Sougb Language-Siwi dialect), ifo-ifo (Tidore Language), nibun kelapa hutan (Malay-Sentani dialect), okiri (Tobaro Language), palindan (Tagalog-Luzon dialect), saser (Wandama language).

uses. Leaves used for thatching, trunk for building houses and bridges, bows or harpoons.

notes. The first mention of this species was by Blanco (1837) in his first edition of the Flora of the Philippines under the name Caryota palindan. Several years later Blanco (1845) produced the second edition of the Flora, citing the same description and adding to it a detailed picture of the species. Unfortunately Blanco did not make herbarium specimens, thus the picture in his second edition has to be regarded as the type of the species (see McNeill et al. 2006). Naves (1883) identified this species as Orania regalis. Although this identification was later shown to be incorrect by Merrill (1905), it was the first time this species was included as a member of the genus Orania. Merrill (1905) transferred Blanco’s Caryota palindan to Orania as O. palindan. As mentioned before, because Blanco did not make specimens, Merrill selected one of his specimens — Species Blancoanae 144 — kept in K as the neotype for O. palindan (see Merrill 1918). The description from this specimen matches Blanco’s original description and picture. Prior to Merrill, Beccari (1885) proposed a new taxon based on a specimen collected from the Bogor Botanic Garden and believed to be of Philippine origin. This species was named O. philippinensis. Beccari (1919a, b) regarded O. palindan as a synonym of O. philippinensis. In the same publication Beccari distinguished one variety, var. sibuyanensis, within ‘O. philippinensis’ on the basis of the characters shown in Table 8. Merrill (1925) regarded O. philippinensis and O. palindan as the same taxon, thus the name O. palindan is the valid name for the Philippine taxon described by Blanco. However, Merrill also recognised the variety, as O. palindan var. sibuyanensis. We have seen only spherical fruits in these taxa and thus do not recognise the varieties. Beccari (1885) described O. moluccana as differing from ‘O. philippinensis’ (i.e. O. palindan) in the shape, size, apex, margin and nerves of the leaflets. Our observations on both the holotype and isotypes of O. moluccana and comparison of these with herbarium specimens of O. palindan indicate that both taxa have elongate-lanceolate leaflets with entire margins and obvious midribs. Length of the leaflets varies from 1 to 2 m and width varies from 4.5 to 7.5 cm in both taxa. In O. palindan the peduncle is generally glabrous in populations in the Philippines and New Guinea, while Moluccan populations (formerly recognised as O. moluccana) have peduncles obviously or densely covered with red-brown tomentum. However, the variation is continuous, so there are several individuals in the Philippines and New Guinea with red-brown tomentum on their peduncles and several individuals have been collected from the Moluccas with rather glabrous peduncles. Actually, Beccari himself considered that the status of O. moluccana as a distinct species was not convincing and that it might one day be placed in synonymy with another species. Thus, O. moluccana is transferred to O. palindan. During observations on spirit-preserved collections we noted that specimens previously known as O. moluccana produce a secretion that changes the clear preservative to dark brown. Nothing is known about this substance and it is not produced by specimens labelled O. palindan. Essig (1980) mentioned that the type specimen of O. moluccana was said to have been collected by Teijsmann & De Vriese from Bacan Island without mentioning any type. Beccari (1885) in the protologue had already written that the first record of the presence of O. moluccana was from a living collection in Bogor Botanic Garden believed to be planted there from either seed or a seedling collected by Teijsmann & De Vriese from an area called ‘Joni’ in Bacan Island, an island within the Moluccas Archipelago (the original annotation text by Beccari himself as follows: ‘Nel Giardino di Buitenzorg si coltiva questa Palma come proveniente da Batcian, dove è chiamata “Joni’ e dove fu trovata dai Sigg. Teijsmann e De Vriese’). Prior to this publication, Beccari (1877) stated that he received a letter from Scheffer (then the Director of Bogor Botanic Garden) mentioning the presence of an Orania from Bacan Island that was said to be identical to O. regalis, as was mentioned by Miquel (1868). Indeed there are two herbarium specimens of this O. moluccana from Bacan Island. The first one is a specimen, kept in L, which was labelled Teijsmann & De Vriese s.n. collected from Bacan Island between 1859 and 1860. The other one is labelled Beccari s.n., kept in FI, collected by Beccari at Bogor Botanic Garden in 1878 — a year after he received the letter from Scheffer — with a note saying of Bacan Island origin (the original annotation text by Beccari himself as follows: ‘Piante del viaggio esequito nell’ anno 1878. Batcian N. Ind. Okur.’). In this study the later specimen (Beccari s.n.) is regarded as the type specimen of O. moluccana for it is likely to be the one used by Beccari. The protologue of O. moluccana itself was published in Ann. Jard. Bot. Buitenzorg in 1885 without mention of the herbarium specimen (Teijsmann & De Vriese s.n.). In this study it is assumed that the living collection that Beccari saw and collected was probably derived from the seed or seedling collected by Teijsmann & De Vriese’s Expedition to Moluccas in 1859 – 1860. We have seen both of the specimens. The specimens collected from Papua record the presence of O. palindan on the mainland of New Guinea, thus making O. palindan the most widespread species in the genus, occurring from Luzon Island in the northern part of the Philippines archipelago — through Sulawesi and the Moluccas — to the mainland of New Guinea. So far the existence of Orania in Sulawesi is only based on a photograph taken by Beckwith (Beckwith 1940 pers. not. see Fairchild 1943) and identified in this study as belonging to O. palindan. Beckwith saw and took a photograph of tall and robust Orania from a mountain (or highland) at approximately 1500 m altitude close to a city called Tanate, which turns out to be in the south-east of Makassar and within the range of Mount Lompobatang. Further fieldwork in Sulawesi is needed. The decision to identify the taxon from Sulawesi as O. palindan, despite the lack of the herbarium specimen is based on comparison with specimens of the same species recently collected from fieldwork in New Guinea. O. palindan in New Guinea is tall and robust: it can reach about 30 to 40 m and diam. of the trunk can be around 40 cm. These make O. palindan the tallest and most robust member of the genus. Fruits of O. palindan collected from individuals planted in Bogor Botanic Garden (previously known as O. moluccana, which here is regarded as a synonym of O. palindan) produce a brown liquid when preserved in alcohol, which colours the alcohol after being kept for one day. The spirit-preserved fruit collections from ‘O. moluccana’ kept in BO and K show the same phenomenon. Fruits from the ‘original’ O. palindan do not produce this liquid, nor do fruits from Papua collected during fieldwork carried out for this study or by others (Baker 2000 pers. comm.). Nothing is known about the nature of this liquid. However, there is no other distinctive morphological character that can be used to separate the individuals that secrete the brown liquid as a distinct taxon.

Table 8 Morphological differences between Orania palindan and O. palindan var. sibuyanensis (Beccari 1919a & 1919b, as O. philippinensis).

6. Orania paraguanensis Becc. (Beccari 1905: 335). Type: O. philippinensis (non Scheff.) Becc.

Orania philippinensis (non Scheff.) Becc. (Beccari 1903: 48). Type: Philippines, Palawan Island, Separation Point, Feb. 1903, Merrill 869 (holotype FI!).

Large palm. Trunk c. 18 – 20 m tall, c. 25 – 30 cm diam. Leaves spirally arranged, c. 3 – 4 m long; rachis with c. 2.4 – 3 cm diam. in the middle, glabrous or rarely with red-brown tomentum; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, 60 – 70 in each side of rachis, leaflets c. 6 cm distant, the proximal leaflets not in a group, the most proximal not the smallest, c. 97 – 150 cm long, 4.5 – 6 cm wide, adaxial surface glabrous, midrib thick, with thin red-brown tomentum, other ribs slender, glabrous, abaxial surface with dense white indumentum, midrib thicker than adaxial, with thin red-brown tomentum, other ribs slender, glabrous. Inflorescence spreading, branching to 2 orders, rarely to 3 orders (Podzorski 765), glabrous or with thin red-brown tomentum, robust; first order branch c. 68 – 70 cm long; rachillae thick, c. 35 – 55 cm long, bearing 63 – 100 flower clusters, bearing triads in the proximal $\frac{1}{5}$ – $\frac{2}{5}$ part, the basal c. 2.5 – 3.5 cm devoid of flowers, triads c. 2 – 2.5 cm distant, rachilla glabrous, with conspicuously pulvinate base. Staminate flowers with calyx of 3 united minute sepals; corolla with 3 free petals, c. 8 mm long, 1.8 – 2 mm wide; stamens always 6, filaments always free, dark brown, c. 0.8 – 0.9 mm long, anthers elongate-lanceolate, pale creamy yellow, 6.5 – 8 mm long; pistillodes absent. Pistillate flowers with calyx of 3 united sepals, c. 1 mm long; corolla of 3 free petals, c. 4 – 6 mm long, 3 – 4 mm wide; staminodes 6, uniform, about ¼ length of gynoecium; gynoecium dark-brown, 2 – 5 mm long, 1.5 – 4 mm wide, stigma of 3 elongate lobes, c. 0.8 – 0.9 mm long. Fruit globose or bilobed, c. 3.6 – 5 cm diam., dull green when young, yellowish green when mature, stigmatic remains sub-basal; epicarp smooth, c. 1 mm thick; mesocarp fibrous, c. 3 – 4 mm thick; endocarp hard, red-brown, c. 0.8 – 1 mm thick; testa thinner than endocarp; endosperm, white or creamy white, c. 2.75 – 3.5 cm diam., with hollow c. 1 cm diam. Embryo placed below the middle line of seed. Eophyll pinnate.

distribution. Palawan Island, Philippines Archipelago and Banguey Island off the coast of Sabah. Moore (1965) mentioned the presence of this species in mainland Sabah itself — somewhere near Marudi Bay — unfortunately no herbarium specimens are known to exist and the record thus is based on Moore’s observation. This species has never been seen in other parts of Borneo. Map 4.

specimens examined. philippines. Palawan: Separation point, Feb. 1903, Merrill 869 (FI! holotype); Taytay, May 1913, Merrill 1316 (BO!, FI! paratype, L!); Taytay, May 1913, Merrill 9269 (FI! paratype, K!); July – Aug. 1925, A. Cenabre FB29998 (BO!, K!); Tangkauarin Area, 5 – 21 March 1984, D. A. Madulid 1017 (K!, L!); Lake Mangueo, Danao, valley of stream leading into NNW bay of lake, 7 April 1984, A. C. Podzorski 765 (K!, L!). MALAYSIA. East Malaysia. Sabah: Banguey Island, July – Sept. 1923, P. Castro & F. Melegrito 1495 (BO!, K!).

habitat. Lowland tropical rainforest.

conservation status. Critically endangered (CR B2a, b). The palm is known from just three localities in Palawan and one in Sabah. We know nothing of its status in Sabah. In Palawan the forest where it grows is subject to intense pressure from mining and land clearance.

etymology. From Paragua, Spanish name for Palawan, an island in the Philippines Archipelago.

vernacular names. banga (Tangkauarin).

uses. Leaves are used for house thatching, ripe fruits are said to be eaten by wild pigs.

notes. One specimen (Podzorski 765) has inflorescence branching to 3 orders; however, the rest of the characters match the description of Orania paraguanensis — including the pinnate eophyll. Prior to this study O. paraguanensis was only known from Palawan. It is now known to occur on Banguey Island (now Banggi Island) just off the coast of Sabah and presumably in Sabah itself — somewhere near Marudi (present day Marudi) Bay (Moore 1965). So now there are two species found in Borneo, O. paraguanensis and O. sylvicola.

The subgroup of species with distichous leaf arrangement

This subgroup consists of the two distichous species from New Guinea, Orania disticha and O. tabubilensis. Despite possession of the same distichous leaf arrangement, O. disticha differs from O. tabubilensis in several characters (Table 9).

Table 9 Morphological differences between Orania disticha and O. tabubilensis.

7. Orania disticha Burret (1935: 321). Type: Papua New Guinea, Central, Kubuna, Nov. 1933, L. J. Brass 5599, (holotype B†, isotypes A!, BO!, NY!).

Robust palm. Trunk c. 15 – 20 m tall, c. 20 – 23 cm diam. (dbh). Leaves 7 – 12 in the crown, distichously arranged, c. 4.3 m long; leaf-sheath and petiole c. 1.3 m long, adaxial surface glabrous, abaxial surface with dense red-brown tomentum, margins disintegrating into fibres; petiole densely covered with red-brown tomentum; rachis c. 3 m long, c. 2.5 – 3 cm diam.; leaflets elongate-lanceolate, regularly arranged, held in one plane, distance between 2 leaflets c. 7 – 8 cm, c. 90 – 150 cm long, 6.5 – 7.5 cm wide, adaxial surface glabrous, shining, with possible wax, midrib robust, other ribs less robust but thick; abaxial surface with white indumentum, thin red-brown tomentum on the margin and some of the ribs, midrib robust. Inflorescence spreading, branching to 2 orders, glabrous, robust, c. 1.2 – 1.3 m long; prophyll persistent, c. 30 × 7 cm, disintegrating into fibres when old; peduncle c. 3 cm diam.; peduncular bract one, woody, persistent, c. 1.3 m long, including c. 15 cm acuminate tip, c. 13 cm wide near base, adaxial surface glabrous, abaxial surface with dense red-brown tomentum, splitting in the middle, disintegrating into fibres when old; first order branches c. 56 – 60 cm long, rachillae bract c. 3 – 5 mm long, 10 mm wide; rachillae robust, conspicuously zigzag, c. 41 – 70 cm long each, bearing 65 – 115 flower clusters, triads proximally arranged up to c. half part of rachilla, distally half part of rachilla with paired staminate flowers, the basal c. 1.5 – 3.5 cm devoid of flowers, triads c. 1.5 – 2.3 cm distant, rachilla glabrous. Staminate flowers pale creamy brown; with calyx of 3 united minute sepals; corolla with 3 free petals, c. 13 – 14 mm long, 4 – 5 mm wide; stamens 6, filaments free, dark brown, c. 1.7 – 2.3 mm long, anthers elongate-lanceolate, pale creamy yellow, always free, c. 6.5 – 7 mm long; pistillode absent. Pistillode flowers pale creamy brown; with calyx of 3 united sepals, c. 2 – 3 mm long; corolla with 3 free petals, c. 8 – 12 mm long, c. 5 – 6 mm wide; staminodes 6, c. 1.3 – 2 mm long; gynoecium dark brown, about 4 × 3 mm; stigma with 3 elongate lobes. Fruit globose or lobed, orange (see Burret 1935), c. 5.5 – 6 cm diam.; epicarp thin, smooth; mesocarp fibrous, c. 3 mm thick; endocarp hard, red brown, c. 1 mm thick; testa very thin, red-brown, covering the endosperm; endosperm homogenous, creamy white, c. 4.5 – 5 cm diam., c. 1.5 cm thick, with hollow inside, c. 2 cm wide. Embryo placed below middle line of seed. Eophyll bifid.

distribution. In Papua this species is found from the tip of the Bird’s Head Peninsula region through Nabire in the central part to somewhere near the Idenburg (now Taritatu) river and the southern part of Papua New Guinea up to the south-central part. Map 6.

Map 6

The distribution of Orania archboldiana (●), O. disticha (■), O. ferruginea (▲), O. macropetala (♦), and O. tabubilensis (▼).

specimens examined. indonesia. Papua Barat. Sorong: Warsamson R., NE of Sorong, 28 Jan. 2002, R. A. Maturbongs RAM 703 (AAU, BO!, K!, LAE). Papua. Nabire: km13 of PT Kaltim Hutama Logging Road, Kwatisore Subdistr., 1 February 2001, R. A. Maturbongs RAM 666 (BO!, K!, MAN). Jayawijaya: Idenburg R., Jan. 1939, L. J. Brass 12407 (BO!, L!). PAPUA NEW GUINEA. Central: Kubuna, Nov. 1933, L. J. Brass 5599 (A!, BO!, NY! isotypi); Veiya, 13 March 1935, Carr 11705 (K!, L!); Veiya, 13 March 1935, Carr 11706 (K!, L!); Kitaki, 9 May 1935, Carr 12232 (BO!; K!); Maipa Airstrip, Kairuku Subprovince, 8 Sept. 1962, Darbyshire 906 (L!). Southern Highlands: near Tage, Lake Kutubu, 21 Sept. 1961, Schodde 2188 (K!, L!).

habitat. Abundant on low ridges of lowland tropical rainforest altitudes about 50 to 100 m and up to margin of primary rainforest in rather higher areas reaching about 810 m above sea level.

conservation status. Near threatened (NT). The palm is known from nine different localities with a wide extent of occurrence but wherever it occurs, the vegetation is subject to extensive logging and disturbance from oil and gas extraction.

etymology. Distichous appearance.

vernacular names. pokengeh (Mekeo-Mapia), tidifa (Kutubu).

uses. Leaves are used for thatching.

notes. In the most recent monograph of the genus (Essig 1980), Orania disticha was the only New Guinean species known to have its leaves arranged distichously. In the present study, we describe another distichous species, O. tabubilensis. Apart from the distichous leaves, O. disticha is very similar to O. palindan. However, we regard distichy as sufficiently important to recognise O. disticha as a species based on recent fieldwork in New Guinea by Baker (2000 pers. comm.) and Maturbongs (2001 pers. comm.). They reported that the distichous leaf arrangement persists in mature individuals found in Madang, Papua New Guinea (Baker 2000 pers. comm.) and Kwatisore-Nabire, Papua (Maturbongs 2001 pers. comm.). Unfortunately, Baker only saw the population from the air on his way to other collecting areas. However, Maturbongs provided us with a good collection. In 2002 Maturbongs collected another specimen of distichous Orania from a different location in Papua, Warsamson area, which is north-east of Sorong in the Bird’s Head Peninsula. They match the descriptions of O. disticha. A claim for another distichous taxon found in New Guinea was proposed by Ferrero (1997). He reported the presence of an individual with a distichous leaf arrangement and colourful leaves in Papua without giving any details of the exact location and suggested it as a new species. In fact the leaf of O. disticha is indeed densely covered with red-brown tomentum and actually in the protologue Burret (1935) clearly noted this. It is the inflorescence, which is glabrous, not the leaf. In this present study it is suggested that until more evidence becomes available, Ferrero’s record is best treated as possibly belonging to O. disticha. The observations on herbarium specimens carried out in this study also agree with the protologue. The other distichous taxa are in Madagascar. Essig (1980) in his monograph wrote that Burret did not designate a holotype and with the Berlin specimen destroyed, Essig selected the specimen in A as lectotype.

8. Orania tabubilensis A. P. Keim & J. Dransf. sp. nov.

Palma parva; foliis distichis dispositis O. distichae simili, sed foliolis irrgulariter dispositis in medio rachidis carentibus differt. Typus: Papua New Guinea, Western, North Fly Distr., road from Tabubil to Ok Tedi copper mine, 8.5 km N of Tabubil, c. 2 km NW of Finalbin, 12 Dec. 2000, W. J. Baker 1129 (holotypus BO!; isotypi AAU, K!, LAE).

http://www.ipni.org/urn:lsid:ipni.org:names:77118908-1

Small palm. Trunk c. 5 m tall, c. 10 cm diam. (DBH). Leaves 8 in crown, arranged distichously, c. 380 cm long, leaf-sheath c. 160 cm long, 18 cm wide near base, adaxial surface glabrous, abaxial surface with dense red-brown tomentum covering, margin disintegrating into fibres, fibres straight; petiole covered with red-brown tomentum, c. 10 – 11 cm long, c. 1.5 – 2 cm diam.; rachis c. 210 cm long, densely covered with red-brown tomentum, c. 1.5 – 2 cm diam.; leaflets elongate-lanceolate, arranged in more than one plane, absent in centre of rachis, c. 35 on either side, c. 90 cm long, 4 – 4.5 cm wide each, distance between 2 leaflets c. 7 cm in near middle part of rachis; adaxial surface glabrous, midrib robust with slight red-brown tomentum covering, other ribs slender; abaxial surface glabrous with slight red-brown tomentum covering on margins, midrib less robust than adaxial surface, other ribs slender. Inflorescence spreading, branching to 2 orders, c. 180 cm long; prophyll c. 52 cm long, distinctly pointed, woody, stiff when dry; peduncle c. 60 cm long, densely covered with red-brown tomentum; peduncular bract apparently one, woody, persistent; rachis c. 120 cm long, densely covered with red-brown tomentum, number of first order branches 25, rachillae rather thick, c. 20 – 50 cm long, slightly zigzag, bearing 42 – 102 flower clusters, bearing triads up to 10 – 25 cm from base-triads arranged in the proximal half part, the basal c. 1 – 1.5 cm devoid of flowers, triads c. 1.5 – 2 cm distant, rachilla densely covered with red-brown tomentum. Staminate flowers with calyx of 3 united sepals, c. 1 mm long; corolla with 3 free petals, c. 6 mm long, 1 – 1.5 mm wide; stamens 6, filaments free, dark-brown, c. 0.5 – 0.75 mm long, anthers elongate-lanceolate, pale creamy yellow, always free, c. 4 mm long; pistillode absent. Pistillate flowers with calyx of 3 minute sepals, 1.5 – 2 mm long; corolla with 3 free petals, c. 2 – 2.5 mm long, 2 mm wide; staminodes 6, c. 0.5 – 1 mm long; gynoecium dark-brown, c. 2 mm long, 2 mm wide, stigma of 3 elongate lobes, 0.5 mm long. Fruit globose, juvenile, c. 2 cm diam., stigmatic remains sub-basal. Embryo placed below middle line of seed. Eophyll unknown. Fig. 1.

distribution. This species possesses an amazingly wide disjunct distribution. In Papua this species is found in Sorong, the most western tip of the Bird’s Head area. In Papua New Guinea the species is found in an area close to a road from Tabubil to Ok Tedi copper mine, 8.5 km N of Tabubil, c. 2 km NW of Finalbin, North Fly Distr., Western Province, Papua New Guinea. Map 6.

specimen examined. indonesia. Papua Barat. Sorong: Km 14 Sorong, 29 January 2002, R. A. Maturbongs RAM 705 (BO!, K!, LAE, MAN). PAPUA NEW GUINEA. Western: North Fly Distr., road from Tabubil to Ok Tedi copper mine, 8.5 km N of Tabubil, c. 2 km NW of Finalbin, S 5°12'31.2' E141°10'42.4', 12 Dec. 2000, W. J. Baker 1129 (holotype BO!, istotypes AAU, K!, LAE).

habitat. In Sorong, Papua this species is found in the lowland forest at altitude about 10 m above sea level. On the contrary, in Papua New Guinea it is found in montane hill forest on limestone soil at about 1000 m above sea level.

conservation status. Vulnerable (VU D2). Although known only from two collections, the palm occurs in some abundance in montane forest on rocky limestone slopes unlikely to be used for agriculture.

etymology. From Tabubil, the type locality.

vernacular names. lai (Sorong, Matbat language).

uses. Unknown.

notes. Prior to this study, Orania archboldiana was the only species in the genus known to have its leaflets arranged in more than one plane (Essig 1980), but that species has spirally arranged leaves. A recently collected specimen from Sorong (R. A. Maturbongs RAM 705) with distichous leaves and irregularly arranged leaflets seems also to belong in O. tabubilensis, despite the astonishingly wide disjunction and difference in habitat preference. The distichous leaf arrangement immediately distinguishes this species from the other species that possesses the same leaflet arrangement, O. deflexa. For a complete list of differences with O. archboldiana and O. deflexa, see Table 12.

The subgroup of species with simply branched inflorescences

Prior to this study Orania parva was the only species known to have its inflorescence branched to one order only; we describe two other simply branched species here for the first time. Differences between the three species are shown in Table 10.

Table 10 Morphological differences between Orania dafonsoroensis, O. timikae and O. parva.

9. Orania dafonsoroensis A. P. Keim & J. Dransf. sp. nov.

Palma grandis; inflorescentia 1-plo ramosa, pedunculo glabro, rachillis semper plus quam 10. Typus: Indonesia, Papua, Cyclops Mountain, near WWF Station Pos 7, Sentani, Jayapura, 18 Feb. 1999, A. P. Keim AK 36 (holotypus K!; isotypi BH!, BO!, MAN!).

http://www.ipni.org/urn:lsid:ipni.org:names:77118909-1

Large palm. Trunk up to 10 m tall, c. 10 – 16 cm diam. (DBH), internodes c. 8 – 10 in middle trunk, 20 cm near the base, dark green coloured. Leaves 10 – 11 in the crown, arranged subdistichously when young, spirally when mature, c. 2.5 – 4.5 m long; leaf-sheath c. 30 – 40 cm long, 9 – 10 cm wide the near base, adaxial surface glabrous, abaxial surface with dense red-brown tomentum covering, margins disintegrating into fibres, fibres slightly curled; petiole covered with red-brown tomentum, c. 70 – 80 cm long, c. 2.3 – 3.5 cm diam.; rachis c. 1.5 – 3.3 m long, covered with red-brown tomentum, c. 2.3 – 2.5 cm diam.; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, c. 70 – 80 in total number, c. 1.03 – 1.70 m long, 8.5 – 9 cm wide each, the proximal part of rachis with single smallest leaflet, distance between 2 leaflets c. 7 cm in middle part of rachis; adaxial surface dark green, glabrous, midrib robust, other ribs slender; abaxial surface densely covered with white indumentum, red-brown tomentum on margin, midrib and near base, midrib robust, other ribs slender but thicker than adaxial. Inflorescence spreading, branching to 1 order, c. 2.8 – 2.94 m long; prophyll persistent, c. 70 – 87.5 cm long, 5 – 6 cm wide, disintegrating into fibres when old; peduncle c. 2 – 2.1 m long, glabrous; peduncular bract one, woody, persistent, c. 3 – 3.1 m long, 5 cm wide, adaxial surface glabrous, abaxial surface slightly to deeply covered with red-brown tomentum, splitting in the middle, disintegrating into fibres when old; rachis c. 80 – 84 cm long, number of rachillae 18 – 20, rachillae thick, c. 45 – 80 cm long, straight, not obviously zigzag, bearing 72 – 128 flower clusters, bearing triads up to 37 – 71 cm from base, the basal c. 4.5 cm devoid of flowers, triads c. 1.5 cm distant, rachilla glabrous. Staminate flowers with calyx of 3 united minute sepals; corolla with 3 free petals, c. 6 × 2 mm; stamens 6, filaments free, dark-brown, c. 0.75 – 1 mm long, anthers elongate-lanceolate, pale creamy yellow, c. 3 mm long; pistillode absent. Pistillate flowers with calyx of 3 united sepals, c. 2 mm long; corolla with 3 free petals, c. 5 × 2 mm; staminodes 6, c. $\frac{1}{3}$ as long as gynoecium, uniform; gynoecium dark-brown, c. 2.5 mm long, 1.5 – 2 mm wide; stigma with 3 lobes. Fruit globose or bilobed, 4 – 4.5 cm diam., dull green when young, bright yellow to orange when mature; epicarp thin, smooth; mesocarp fibrous, c. 2 – 3 mm thick; endocarp thin, dark-brown, hard; endosperm white, c. 3.5 – 4 cm diam., with hollow inside, liquid endosperm present. Embryo placed below middle line of seed. Eophyll bifid. Fig. 2.

distribution. New Guinea. Endemic in the Cyclops (Dafonsoro in local Sentani language) mountain in northern part of Papua around the Sentani Lake and close to Jayapura. Map 7.

Map 7

The distribution of Orania dafonsoroensis (♦), O. timikae (▼), O. littoralis (▲), O. oreophila (■), O. parva (◊), O. regalis (●), and O. subdisticha (□).

specimens examined. indonesia. Papua. Jayapura: Cyclops Mountain, 12 Aug. 1998, C. Heatubun CH 278 (K!, MAN!); near WWF Station at Pos 7-Sentani, 18 Feb. 1999, A. P. Keim AK 35 (BH!, BO!, K!, MAN!); A. P. Keim AK 36 (BH!, BO!, K! holotype, MAN!); North Cyclops, 1 Feb. 2001, B. Desianto BD 12 (AAU, CANB, K!, MAN).

habitat. Mountain slope in undisturbed humid rather hilly tropical rainforest with altitude from 700 up to 900 m above sea level.

conservation status. Critically endangered (CR). The palm occurs in a single locality, which, despite being within a national park, is vulnerable to continuing disturbance and degradation.

etymology. From Dafonsoro, an indigenous name for Cyclops Mountain where the type was collected.

vernacular names. hara cho (Sentani, northern Cyclops).

uses. Unknown.

notes. This species is one of three species with simply branching inflorescences. The existence of this species was recognised in 1998 by a team of researchers from the National University of Papua (then Cenderawasih University at Manokwari), when one collection was made (C. Heatubun CH 278). Another collection was made in 2001 in the northern part of the mountain (B. Desianto BD 12). This is a beautiful palm with an outstanding long peduncle and big fruits hanging on the rachillae. Fortunately, the Cyclops Mountain is a Nature Reserve (Schultz-Westrum 1978; Nasution et al. 1995) and thus this species should survive.

10. Orania timikae A. P. Keim & J. Dransf. sp. nov.

Palma parva; folia subdisticha disposita; inflorescentia 1-plo ramosa, pedunculus cum densus tectus rubellus brunneus tomentosus, rachillis 10. Typus: Indonesia, Papua, Fakfak, path E at Mile 50 on the road to Tembagapura, Timika, 11 Feb. 1998, J. Dransfield JD 7667 (holotypus K!; isotypi BO, MAN).

http://www.ipni.org/urn:lsid:ipni.org:names:77118910-1

Small palm. Trunk 4 m tall, c. 10 cm diam. (dbh), internodes c. 1.5 – 2 cm. Leaves 6 in crown, subdistichously arranged, c. 3 m each long; petiole c. 2 cm diam., with dense red-brown tomentum; rachis c. 1.5 – 2 cm diam., with dense red-brown tomentum; leaflets regularly arranged — held in one plane, 23 leaflets on each side of rachis, c. 88 – 90 cm long, 5 – 6 cm wide, leaflets c. 10 cm distant, adaxial surface green, glabrous, with red-brown tomentum on the midrib and some of the other ribs, midrib thick, others slender, abaxial surface densely covered with white indumentum, red-brown tomentum on every rib and conspicuously on the base of leaflets, midrib slender. Inflorescence spreading, branching to 1 order, c. 150 cm long, with dense red-brown tomentum; prophyll persistent, c. 25 cm long, 3.5 cm wide, adaxial surface glabrous, abaxial surface with red-brown tomentum; peduncle c. 100 cm long, glabrous; peduncular bract one, c. 150 – 160 cm long, 3 – 3.8 cm wide, persistent, woody, adaxial surface glabrous, abaxial surface densely covered with red-brown tomentum, splitting in the middle, from top to the base, heavily disintegrating into fibres when old; rachis c. 50 cm long; rachillae slender, 10, c. 44 – 45 cm long each, bearing 80 – 85 flower clusters, bearing triads arranged in the proximal half part, the basal c. 3.5 cm devoid of flowers, triads c. 1 – 1.5 cm distant, rachilla glabrous. Staminate flowers reddish dark brown, with calyx of 3 united sepals, c. 1 – 1.5 mm long; corolla with 3 free petals, c. 8 mm long, 2.5 – 3 mm wide; stamens 6, 6 mm long, filaments always free, uniform, dark-brown, about 1 mm long, anthers elongate-lanceolate, pale yellow, always free, uniform, c. 5 mm long; pistillodes absent. Pistillate flowers reddish dark brown, with calyx of 3 sepals, c. 2.5 – 3 mm long; corolla with 3 free petals, c. 6 mm long, 4 mm wide; staminodes 6, uniform, c. 0.8 – 1 mm long; gynoecium dark brown, c. 4 – 5 mm long, 2 – 3 mm wide, stigma with 3 elongate lobes, 2 – 2.5 mm long. Fruits unknown. Embryo unknown. Eophyll apparently bifid (Dransfield 2003 pers. comm.). Fig. 3.

distribution. Endemic to an area between Timika and Tembagapura in the vicinity of a copper mine concession in Papua. Map 7.

specimens examined. indonesia. Papua. Fakfak: km 63 on road to Tembagapura Loc.1, 04°20'44″S 136°58'12''E, 7 Feb. 1998. C. Heatubun CH 173 (BO!, BH!, K!, L!, MAN!); path E at Mile 50 on road to Tembagapura, Timika, 4°17'14''S 137°1'1''E, 11 Feb. 1998, J. Dransfield JD 7667 (BO!, K! holotype, MAN).

habitat. Orania timikae grows on a slope in forest transitional between lowland rainforest and heath forest at about 540 m altitude Other populations were seen and collected in the middle of a heath forest at about 435 m altitude.

conservation status. Vulnerable (VU D2). Although the palm is known from a restricted area, the local vegetation is heath forest, unsuitable for agriculture, and receives some protection by virtue of its being within the PT Freeport concession.

etymology. From Timika, the type locality.

vernacular name. Unknown.

uses. Unknown.

notes. This species was found during fieldwork in 1998. It has been collected from a rather unusual and unlikely habitat for Orania — heath forest. So far, only O. sylvicola in Sumatra and Borneo is known to grow in this type of forest. In the field O. timikae may look very similar to either O. dafonsoroensis or O. parva, but as well as the unusual habitat, O. timikae is distinct because it is the only member of the subgroup that possesses the subdistichous leaf arrangement, which persists through maturity. O. dafonsoroensis has the subdistichous leaf arrangement only at the juvenile stage. O. parva always has spiral leaf arrangement. Other distinguishing characters are shown in Table 10. As for O. dafonsoroensis, this species may have a potential use as an ornamental palm.

11. Orania parva Essig (1980: 226). Type: Papua New Guinea, West Sepik, Carpentaria Exploration Co., Frieda R., Telefomin, 26 April 1978, Essig & Young LAE 74046 (holotype LAE; isotype BH!).

Small palm. Trunk 4 m tall, c. 8 cm diam. (dbh). Leaves spirally arranged, 200 – 204 cm long; petiole c. 2 cm diam., with dense red-brown tomentum; rachis c. 2 cm diam., with dense red-brown tomentum; leaflets regularly arranged, held in one plane, 23 leaflets on each side of rachis, c. 90 – 106 cm long, 5.5 – 6 cm wide, c. 8 – 13 cm distant, adaxial surface green, glabrous, with red-brown tomentum on the midrib and some of the other ribs, midrib thick, others slender, abaxial surface densely covered with white indumentum, red-brown tomentum on every rib and conspicuously on the base of leaflets, midrib slender. Inflorescence spreading, branching to 1 order, c. 1.18 m (118 cm) long; prophyll persistent, c. 38.5 cm long, 3.5 – 3.6 cm wide, adaxial surface glabrous, abaxial surface with red-brown tomentum; peduncle c. 81 cm long, glabrous; peduncular bract one, c. 128 – 130 cm long, 3 – 3.8 cm wide, persistent, woody, adaxial surface glabrous, abaxial surface densely covered with red-brown tomentum, splitting in the middle, from top to the base, heavily disintegrating into fibres when old; rachis c. 37 cm long; rachillae slender, 8 – 10, c. 35 – 36 cm long each, bearing 120 – 128 flower clusters, bearing triads arranged in the proximal half part, the basal c. 3.5 cm devoid of flowers, triads c. 1 – 1.5 cm distant, rachilla glabrous. Staminate flowers creamy white, with calyx of 3 united sepals, c. 2 – 3 mm long; corolla with 3 free petals, c. 5 – 8 mm long, 2 mm wide; stamens 6, 5 mm long, filaments always free, dark-brown, about 2 mm long, anthers elongate-lanceolate, pale yellow, c. 3 mm long; pistillodes absent. Pistillate flowers creamy white, with calyx of 3 sepals, c. 2 – 2.5 mm long; corolla with 3 free petals, c. 6 – 7 mm long, 3 – 4 mm wide; staminodes 6, uniform, c. 0.7 – 1 mm long; gynoecium dark brown, c. 2 – 4 mm long, 2 – 3 mm wide, stigma with 3 elongate lobes. Fruits globose or bilobed, c. 1.3 cm diam. when young. Embryo placed below middle line of seed. Eophyll bifid.

distribution. New Guinea. In Papua New Guinea known only from the type locality, with a number of individuals seen (Essig 1980). Map 7.

specimens examined. papua new guinea. West Sepik: Carpentaria Exploration Co., Frieda R., Telefomin, 4°40'S 139°55'E, 26 April 1978, Essig & Young LAE 74046 (BH!, LAE holotype).

habitat. Orania parva grows on well-drained slopes at about 60 m above sea level.

conservation status. Data deficient (DD). The palm is known only from its type and we have no recent information on the status of forests in the type locality.

etymology. From Latin, parvus means small.

vernacular name. Unknown.

uses. Unknown.

notes. Orania parva is the smallest member of the genus. This is a beautiful small palm with a simply branching inflorescence and a very long peduncle. Essig (1980) mentioned that it often flowers within arm’s reach and although only the type collection was made, a number of individuals were seen in the surrounding area and agreed well with the type. Table 10 shows the differences between this species and the other two members of the subgroup.

The subgroup of species with small-understorey habit and inflorescences branched to more than one order

Five species belong to this subgroup (Orania archboldiana, O. ferruginea, O. littoralis, O. oreophila and O. subdisticha). The differences between the species are listed in Table 11.

Table 11 Features distinguishing Orania archboldiana, O. ferruginea, O. littoralis, O. oreophila, and O. parva.

12. Orania archboldiana Burret (1939: 198). Type: Papua New Guinea, Western, Lower Fly R., east bank opposite Stuart Island, Oct. 1936, L. J. Brass 8225 (lectotype A!, selected here).

Small to medium palm. Trunk up to 15 m tall, c. 6 – 10 cm diam. near the crown. Leaves 6 – 10 in the crown, spirally arranged, c. 2.7 – 3 m long; leaf-sheath c. 22 – 30 cm long, 5 cm wide, adaxial surface glabrous, abaxial surface densely covered with red-brown tomentum, margin disintegrating into fibres; petiole 90 cm long; lamina c. 1.6 – 2.25 m long, in middle part c. 1.5 – 2 cm diam.; leaflets elongate-lanceolate, regularly arranged, but held in more than one plane giving the whole leaf a plumose appearance, the proximal 3 leaflets crowded 1.5 – 2 cm distant in a group, otherwise c. 4.5 cm distant in middle part of rachis, distance between 2 leaflets in the proximal part c. 4.5 cm, otherwise c. 1.5 – 2 cm distant in middle part of rachis, leaflets c. 60 – 90 cm long, 3 – 4.7 cm wide, adaxial surface glabrous, except for sparse red-brown tomentum on the midrib, midrib thick, other ribs slender, glabrous, abaxial surface densely covered with white indumentum, red-brown tomentum on the margin and midrib, midrib massive, other ribs glabrous and slender. Inflorescence spreading, branching to 2 orders, c. 90 – 133 cm long; prophyll persistent, disintegrating into fibres from top to base; peduncle c. 30 – 45 cm long, covered with sparse red-brown tomentum; peduncular bract 1, woody, splitting in the middle, persistent, abaxially with sparse red-brown tomentum; rachis 60 – 88 cm long; first order branches c. 60 – 70 cm long; rachillae slender, straight, not conspicuously zigzag, c. 30 – 50 cm long, bearing 90 – 153 flower clusters, bearing distichously or subdistichously arranged triads in the proximal $\frac{3}{4}$ and paired staminate flowers in the distal $\frac{1}{4}$, the basal c. 0.7 – 2.5 cm devoid of flowers, triads c. 6 – 10 mm distant, rachilla surface glabrous. Staminate flowers with calyx of 3 united minute, scale-like sepals; corolla with 3 free petals, c. 4 × 1 mm; stamens 6, filaments free, dark brown, c. 0.75 mm long, anthers elongate-lanceolate, pale creamy-yellow, always free, c. 3 mm long each; pistillodes absent. Pistillate flowers with calyx of 3 minute united sepals; corolla with 3 free petals, c. 2 to 3 mm long, 1 – 2 mm wide; staminodes 6, c. $\frac{1}{3}$ – $\frac{1}{2}$ as long as the gynoecium; gynoecium dark brown, c. 1 – 1.5 mm long; stigma of 3 elongate lobes. Fruits globose or bilobed, c. 3.5 cm diam. Embryo placed below middle line of seed. Eophyll bifid.

distribution. Southern part of New Guinea, both Papua and Papua New Guinea. In Papua Orania archboldiana so far has only been collected from an area around Merauke, close to the border with the Western province of Papua New Guinea. In Papua New Guinea the species is found scattered in an area about latitude 6°10'S to 8°55'S and longitude 141°20'E to 144°13'E. Map 6.

specimens examined. indonesia. Papua. Merauke: Kwell village, 2 Oct. 2000, R. A. Maturbongs RAM 655 (AAU, BO!, K!, MAN); R. A. Maturbongs RAM 656 (AAU, BO!, K!, MAN). PAPUA NEW GUINEA. Western: Oroville Camp–Fly R., Aug. 1936, L. J. Brass 7403 (A! Paratype); Lower Fly R., E bank opposite Stuart Island, Oct. 1936, L. J. Brass 8184 (A! Paratype); L. J. Brass 8225 (A! lectotype); Kiunga-Kiunga Subprovince, 10 Aug. 1971, Streimann & Katik LAE 51793 (K!, L!, LAE); Pawa Village on Oriomo R., Daru Subprovince, 16 July 1974, Foreman & Stocker LAE 60431 (L!). Gulf: Kikori distr., Kopi – Kikori road, 3 km NW of Kikori, 7°24'12.4''S, 144°13'56.2''E, 22 Nov. 2000, W. J. Baker 1104 (BO!, K!, LAE, AAU, NY, L).

habitat. On ridges in lowland tropical rainforest about 20 m above sea level.

conservation status. Near Threatened (NT). The palm is known from seven different localities with a wide extent of occurrence but wherever it occurs, the vegetation is subject to extensive logging and disturbance from oil and gas extraction.

etymology. After Richard Archbold, an entrepreneur who financed several botanical expeditions.

vernacular name. Unknown.

uses. Unknown.

notes. Prior to this publication (Essig 1980) Orania archboldiana was the only species known to have its leaflets arranged in more than one plane. Differences between it, O. tabubilensis and O. deflexa are shown in Table 12.

Table 12 Morphological features distinguishing Orania archboldiana, O. deflexa, and O. tabubilensis.

13. Orania ferruginea A. P. Keim & J. Dransf. sp. nov.

Palma parva; a ceteris speciebus parvis Oraniae inflorescentia 2-plo ramosa 100 ad 110 fasciculis florumper rachilla distincta. Typus: Indonesia, Papua Barat, Manokwari, Tuwanwowi, protected forest of ‘Balai Latihan Kehutanan Nasional’ (National Educational Forestry Bureau), 22 March 1999, A. P. Keim AK 42 (holotypus K!; isotypi BO!, MAN!).

http://www.ipni.org/urn:lsid:ipni.org:names:77118911-1

Small palm. Trunk c. 10 m tall, c. 5 – 6 cm diam. (dbh); internodes green, smooth, c. 10 cm. Leaves 10 in a crown, regularly arranged, c. 2.3 – 4 m long; leaf-sheath c. 40 cm long, c. 9 cm wide near base, margins disintegrating into fibres, fibres straight, c. 10 – 12 cm long, adaxial surface glabrous, abaxial surface with dense creamy white indumentum and red-brown tomentum; petiole c. 80 cm long, c. 2 cm diam., densely covered with red-brown tomentum and creamy white indumentum; rachis c. 1.1 – 2.8 m long, c. 2 cm diam. in the middle part, densely covered with red-brown tomentum and creamy white indumentum; leaflets elongate-lanceolate, regularly arranged leaflets held one plane, leaflets c. 6 – 7 cm distant, c. 87 – 100 cm long, 5 cm wide, adaxial surface glabrous, red-brown tomentum on midrib, midrib thick, other ribs slender, abaxial surface with dense white indumentum, red-brown tomentum on midrib and margin, midrib thick, other ribs slender. Inflorescence spreading, branching to 2 orders, c. 1 – 1.2 m long, slender; prophyll persistent, c. 17 – 25 cm long, 4 – 4.5 cm wide near base, margins disintegrating into fibres, fibres straight, c. 10 – 12 cm long; peduncle c. 50 – 56 cm long, deeply covered with red-brown tomentum; peduncular bract one, woody, c. 1.14 – 1.4 m long, 6 – 7.5 cm wide in near base, persistent, splitting in the middle, disintegrating into fibres when old, adaxial surface glabrous, abaxial surface with dense red-brown tomentum; rachis c. 50 – 64 cm long; first order branches c. 24 – 25 cm long, rachillae bract small; rachillae slender, c. 18.5 – 20 cm long, bearing 100 – 150 flower clusters, arranged in triads in the proximal $\frac{1}{3}$, the basal c. 1 – 2 cm devoid of flowers, triads c. 1 cm distant. Flowers unknown. Fruits globose or bilobed, c. 3.5 – 4 cm diam., dull green when young, yellow when mature. Embryo placed below middle line of seed. Eophyll bifid. Fig. 4.

distribution. New Guinea. Within the mainland of New Guinea this species has a disjunct distribution, found in an area called Tuwanwowi near the town of Manokwari in Papua Barat and a hill called Bisidak near the town of Elelim in the Kurima Distr. in Wamena vicinity, Papua. Map 6.

specimens examined. indonesia. Papua Barat. Manokwari: Tuwanwowi, protected forest of BLKN, 22 March 1999, A. P. Keim AK 41 (BO!, K!, MAN!); A. P. Keim AK 42 (BO!, K! holotype, MAN!). Papua. Wamena: Kurima subdistr., Elelim, Bukit Bisidak, 11 Oct. 1993, D. Gandawidjaja 2369 (BO!).

habitat. Lowland undisturbed tropical rainforest with altitudes about 200 up to 600 m.

conservation status. Critically endangered (CR B1, B2, C). The palm occurs in two localities. The type locality near Manokwari is within a protected area but the population size is very restricted. We know little of the second locality near Wamena.

etymology. Rusty red-brown coloured.

vernacular names. Unknown.

uses. Unknown.

notes. Orania ferruginea is easily recognised by its dense and thick red-brown tomentum covering on the leaf and infructescences.

14. Orania littoralis A. P. Keim & J. Dransf. sp. nov.

Palma parva; a cateris speciebus parvis Oraniae inflorescentia 2-plo ramosa 70 ad 78 fasciculis florumper rachilla distincta; inventus ad ora restrictus vel littoralis. Typus: Papua New Guinea, Milne Bay, end of logging road, 2 March 2000, Barford et al. 456 (holotypus K!; isotypi AAU, BRI, CANB, LAE).

http://www.ipni.org/urn:lsid:ipni.org:names:77118912-1

Small palm. Trunk 6 m tall, c. 14 cm diam. (dbh), internodes 5 – 5.5 cm, leaf scars inconspicuous basally, visible on upper part, 3.5 – 4 cm wide below crown. Leaves 12 in crown, spirally arranged, c. 6 m long each; leaf-sheath c. 2 m long, margin disintegrating into fibres, adaxial surface glabrous, abaxial surface with dense red-brown tomentum; petiole short, c. 2 cm diam., with dense red-brown tomentum; rachis c. 4 m long, c. 2 cm diam., with dense red-brown tomentum; leaflets elongate-lanceolate, regularly-arranged leaflets held in one plane, leaflets c. 8 cm distant in the middle part of rachis, c. 67 leaflets on each side of rachis, c. 120 cm long, c. 7 cm wide, adaxial surface bright green, shiny, main vein obvious, other veins slender, abaxial surface with dense white indumentum and red-brown tomentum on main vein, other veins slender, glabrous. Inflorescence spreading, branching to 2 orders, c. 245 cm long; prophyll persistent, c. 63 cm long, disintegrating into fibres when old, adaxial surface greenish yellow, glabrous, abaxial surface with red-brown tomentum; peduncle c. 85 cm long, c. 7 cm diam., with dense red-brown tomentum, c. 3.5 cm diam.; peduncular bract one, woody, c. 240 cm long, acuminate tip, c. 42 cm long, adaxial surface glabrous, abaxial surface with dense red-brown tomentum, splitting in the middle, disintegrating into fibres; rachis c. 160 cm long, with dense red-brown tomentum; first order branches 23, c. 63 – 80 cm each; rachillae thick, not robust, straight, not conspicuously zigzag, c. 40 – 65 cm long, bearing 70 – 78 flower clusters, triads arranged in the half part, c. 2 cm devoid of flowers, triads c. 2 – 2.5 cm distant, rachilla with sparse red-brown tomentum. Staminate flowers with calyx of 3 united sepals, c. 1 mm long; corolla with 3 free petals, c. 9 – 10 mm long, 2 mm wide; stamens 6, filaments free, dark-brown, c. 1 mm long, anthers elongate-lanceolate, pale creamy yellow, always free, c. 6 – 7 mm long; pistillode absent. Pistillate flowers with calyx of 3 united sepals, 1 mm long; corolla with 3 free petals, c. 4 mm long, 3.5 – 4 mm wide; staminodes 6, c. 2 mm long, uniform, tips not hooked; gynoecium dark blackish brown, c. 3 mm long, 4 mm wide, stigma of 3 elongate lobes, 0.8 – 0.9 mm long. Fruit globose or bilobed, c. 4.5 – 5 cm diam., stigmatic remains sub-basal, mature orange to red; endosperm white or creamy white, c. 3 – 3.5 cm diam. Embryo placed below middle line of seed. Eophyll bifid. Fig. 5.

distribution. New Guinea. Endemic to an area within the Milne Bay Province, Papua New Guinea. Map 7.

specimen examined. papua new guinea. Milne Bay: end of logging road, 10°22'S 150°7'E, 2 March 2000, Barfod et al. 456 (AAU, BRI, CANB, K! holotype, LAE).

habitat. Lowland rainforest at about sea level.

conservation status. Critically endangered (CR). The palm is known only from its type, collected at the end of a logging road. It seems highly likely that the palm is threatened with further logging in the area.

etymology. Seacoast-loving.

vernacular names. Unknown.

uses. Unknown.

notes. Orania littoralis shares many similarities with both O. ferruginea and O. oreophila. However, several morphological characters can be used to distinguish them (Table 11). In terms of habitat preferences, O. littoralis differs from O. oreophila in growing in lowland forest at sea-level compared with the montane O. oreophila.

15. Orania oreophila Essig (1980: 225 – 226). Type: Papua New Guinea, Eastern Highlands, Kainantu Sub-province, Saboa, Anandara Village, 20 Jan. 1972, Essig LAE 55147 (holotype BH!; isotype LAE).

Small palm. Trunk c. 5 cm diam. (dbh). Leaves 5 – 6 in the crown, spirally arranged, c. 275 – 310 cm long; leaf-sheath c. 20 cm long, margins disintegrating into fibres, adaxial surface glabrous, abaxial surface with dense red-brown tomentum; petiole 100 – 120 cm long, with thin red-brown tomentum; rachis 155 to 170 cm long, c. 1 cm diam. in the middle, with dense red-brown tomentum; leaflets elongate-lanceolate, spirally arranged leaflets held in one plane, the proximal 2 leaflets crowded in a group, otherwise c. 7 – 8 cm distant in the middle, 22 – 24 on each side, c. 60 – 78 cm long, 4.5 – 5 cm wide, adaxial surface dark green, glabrous, with red-brown tomentum on midrib, midrib thick, other ribs slender, glabrous, abaxial surface with white indumentum, red-brown tomentum on the margin, ribs slender. Inflorescence spreading, branching to 2 orders, covered with red-brown tomentum, no sign of wax; prophyll persistent, c. 26.5 cm long, 4.0 – 4.5 cm wide, adaxial surface glabrous, abaxial surface with thin red-brown tomentum, disintegrating into fibres when old, from upper part down to the base; peduncular bract one, woody, c. 58.5 – 59 cm long, 1.6 – 1.7 cm wide, adaxial surface glabrous, abaxial surface with red-brown tomentum, absent in the old inflorescence; first order branch c. 28 – 30 cm long, rachilla bract c. 3 mm long, rachillae slender, c. 18 – 24 cm long, bearing triads in the proximal $\frac{4}{5}$ part and paired staminate flowers in the distal $\frac{1}{5}$ part, the basal c. 1 – 1.5 cm devoid of flowers, triads c. 1 – 1.5 cm distant, rachilla glabrous, bearing 48 to 67 flower clusters, the proximal 22 – 37 contains pistillate flowers. Staminate flowers with calyx of 3 united minute sepals; corolla with 3 free petals, c. 8 mm long, 2.5 – 3 mm wide; stamens 6, filaments free, dark brown, c. 1 mm long, anthers elongate-lanceolate, pale creamy yellow, c. 3.5 – 4 mm long; pistillodes absent. Pistillate flowers with calyx of 3 united minute sepals; corolla with 3 free petals, c. 5 × 3 mm; staminodes 6, c. 1 mm long, uniform; gynoecium dark brown, c. 3 mm long, stigma with 3 elongate lobes, c. 0.75 mm long. Fruits globose or bilobed, c. 3 – 4.5 cm diam. Embryo placed below middle line of seed. Eophyll bifid.

distribution. Papua New Guinea: mountains of the Kainantu Sub-province and Bismarck Mountains in Eastern Highlands Province to mountains of the western part of Morobe Province. Always at high elevation within a range of approx. 1281 – 1620 m (4200 – 5300 feet). Map 7.

specimens examined. papua new guinea. Eastern Highlands: Kainantu Sub-province, Saboa, Anandara Village, 20 Jan. 1972, Essig LAE 55147 (BH! holotype, LAE); Kassam Pass, 11 Jan. 1968, Henty & Coode NGF 29245 (L!).

habitat. In Castanopsis dominated highland forest. Recorded from 6°12'S to 6°28'S latitude and 146°02'E to 146°05'E longitude.

conservation status. Data deficient (DD). The palm is known from only two specimens and we have no recent information on the status of forests at these localities.

etymology. Mountain-loving.

vernacular names. Unknown.

uses. Unknown.

notes. Orania oreophila is one of the smaller members of the genus. It occupies the highest habitat known in the genus. Essig (1980) mentioned the specimen cited in his monograph (Essig LAE 55142), which has an unusual appearance in having large, well-developed bracts on the upper peduncle and subtending the branches. The significance of these characters is uncertain because none of the other specimens has these features.

16. Orania subdisticha A. P. Keim & J. Dransf. sp. nov.

Palma parva, folia sub-disticha disposita; inflorescentia 2-plo ramose. Typus: Papua New Guinea, Southern Highlands, Erave Distr., road to main Gobe oil rig, 91 km NW of Kikori, 26 Nov. 2000, W. J. Baker 1116 (holotypus BO!; isotypi AAU, K!, L, LAE, NY).

http://www.ipni.org/urn:lsid:ipni.org:names:77118913-1

Small palm. Trunk c. 8 m tall, c. 8 cm diam. (DBH). Leaves 7 in the crown, arranged sub-distichously, c. 300 cm long; leaf-sheath c. 120 cm long, 14 cm wide in the near base, adaxial surface glabrous, abaxial surface with dense red-brown tomentum covering, margins slightly disintegrating into fibres; petiole covered with red-brown tomentum, c. 26 cm long, c. 2 – 2.5 cm diam.; rachis c. 154 cm long, densely covered with red-brown tomentum, c. 2 cm diam.; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, c. 22 on either side, c. 70.5 – 80 cm long, 5 – 5.5 cm wide, distance between 2 leaflets c. 10 cm in middle part of rachis; adaxial surface dark green, midrib robust with red-brown tomentum covering, other ribs slender; abaxial surface densely covered with white indumentum, red-brown tomentum on ribs, margin, midrib and near base, midrib less robust than adaxial surface, other ribs slender but thicker than adaxial. Inflorescence spreading, branching to 2 orders, c. 120 cm long; prophyll unknown; peduncle c. 60 cm long, apparently glabrous; peduncular bract apparently one, woody, persistent, c. 150 cm long; rachis c. 60 cm long, number of first order branches 13, rachillae slender, c. 33 – 50 cm long, straight, not obviously zigzag, bearing 63 – 90 flower clusters, bearing triads up to 15 – 25 cm from base (i.e. triads arranged in the proximal $\frac{1}{3}$), the basal c. 1 cm devoid of flowers, triads c. 1 cm distant, rachilla glabrous. Staminate flowers with green petals, remaining characters unknown. Pistillate flowers with calyx of 3 united sepals, c. 2 mm long; corolla with 3 free yellowish green, c. 7 × 5 mm; staminodes 6, c. $\frac{1}{3}$ as long as gynoecium, uniform; gynoecium dark-brown, c. 4 – 5 mm long, 4 – 5 mm wide; stigma with 3 lobes. Fruit globose, c. 4 cm diam., red. Embryo unknown. Eophyll unknown. Fig. 6.

distribution. New Guinea. Endemic to an area close to a road to main Gobe Oil Rig, 91 km NW of Kikori, Erave Distr., Southern Highlands. Map 7.

specimen examined. papua new guinea. Southern Highlands: Erave Distr., road to main Gobe oil rig, 91 km NW of Kikori, 6°46'45.7''S 143°43'59''E, 26 Nov. 2000, W. J. Baker 1116 (AAU, holotype BO!, K!, L, LAE, NY).

habitat. Montane forest on karst limestone with numerous cliffs and sinkholes at an altitude about 1050 m above sea level.

conservation status. Vulnerable (VU D2). Although known only from a single collection, the palm occurs in some abundance in montane forest on rocky limestone slopes unlikely to be used for agriculture.

etymology. Sub-distichous leaf arrangement.

vernacular names. Unknown.

uses. Unknown.

notes. Although the subdistichous leaf arrangement is also observed in Orania dafonsoroensis and O. timikae, O. subdisticha has its inflorescence branched to two orders, whereas O. dafonsoroensis and O. timikae have their inflorescence branched to one order only (i.e. simply branched). For other differences between the species see Table 13. As leaf arrangement in Orania may change with maturity, the question arises as to whether O. subdisticha may just be a subdistichous form of O. disticha. We have found that apart from the subdistichous leaves, O. subdisticha differs from O. disticha in several morphological characters. O. subdisticha is a small understorey palm, whereas O. disticha is a robust palm with trunk from 15 to 20 m tall and diam. from 20 to 23 cm. O. subdisticha apparently possesses slender inflorescences and straight rachillae, whereas in O. disticha they are both robust and have conspicuously zigzag rachillae. The fruits are also different in size (as measured by their diam.) — smaller in O. subdisticha (about 4 cm compared to 5.5 – 6 cm in O. disticha).

Table 13 Morphological features distinguishing Orania timikae and O. subdisticha.

The subgroup ‘Lauterbachiana complex’

Differences between Orania lauterbachiana and O. micrantha are shown in Table 14. The results of this study agree with Beccari (1915) in recognising O. lauterbachiana and O. micrantha as distinct species (O. brassii and O. clemensiae were unknown to Beccari). In contrast, the results disagree with Burret (1936) in recognising O. brassii, O. clemensiae, O. lauterbachiana and O. micrantha as distinct from one another. There are no substantial differences between O. lauterbachiana and O. brassii, or between O. micrantha and O. clemensiae (Table 15). The results of this study are in accordance with Essig (1980) in placing O. brassii as synonym of O. lauterbachiana, but disagree in placing O. micrantha as a synonym of O. lauterbachiana.

Table 14 Comparison between members of ‘Lauterbachiana complex’.

Table 15 Comparison between Orania brassii, O. clemensiae, O. lauterbachiana, and O. micrantha.

17. Orania lauterbachiana Becc. (Beccari 1915: 36). Type: Papua New Guinea, North East, Gogol Mittelauf, 17 Nov. 1890, Lauterbach 970 (holotype B†; isotype FI!).

Orania brassii Burret (1936: 68). Type: Papua New Guinea, Central Province, Mafulu, 12 Nov. 1933, L. J. Brass 5489 (lectotype NY!, selected here; isolectotypes B†, BO!), synon. nov.

Large palm. Trunk up to 10 – 20 m tall, c. 18 cm diam. near the crown, c. 32 cm diam. near the base. Leaves 10 in the crown, spirally arranged, c. 3.5 – 5.1 m long; Leaf-sheath c. 30 cm long, margins disintegrating into fibres, c. 13 – 15 cm long, adaxial surface glabrous, abaxial surface with red-brown tomentum; petiole c. 1.2 – 2.20 m long, covered with red-brown tomentum in the proximal part, c. 2.3 – 2.4 cm diam.; rachis densely covered with red-brown tomentum, c. 2 cm diam. in the middle part; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, 18 – 21 on each side of rachis, sometimes the proximal 2 leaflets crowded in a group (L. J. Brass 5489), otherwise c. 5.5 – 15 cm distant, c. 70 – 132 cm long, 5.5 – 11.5 cm wide, adaxial surface glabrous or with thin red-brown tomentum, mainly near the base, with white appearance, midrib slender, other ribs more slender, abaxial surface with dense white indumentum, red-brown tomentum on margin, midrib thick, other ribs more slender. Inflorescence spreading, branching into 2 orders, c. 95 – 192 cm long, robust; prophyll persistent, hard, splitting in the middle, c. 45 – 161 cm long, 7 cm wide, disintegrating into fibres when old; peduncle c. 53 – 65 cm long, with thin or dense red-brown tomentum; peduncular bract division unknown; rachis c. 57 – 127 cm long; first order branches c. 50 – 90 cm long; rachillae thick, rarely slender, distally zigzagging, with thin red-brown tomentum, c. 35 – 60 cm long, bearing 96 – 104 flower clusters, bearing triads arranged in the proximal $\frac{1}{2}$ – rarely in the proximal $\frac{2}{5}$ & paired staminate flowers in the distal $\frac{3}{5}$ part, the basal c. 2 – 3.5 cm devoid of flowers, triads c. 1.5 – 2 cm distant, rarely 0.5 – 1 cm distant, rachilla glabrous or with thin red-brown tomentum, some deeply textured. Staminate flowers pale creamy white; with calyx of 3 united sepals, c. 1 – 2 mm long; corolla with 3 free petals, c. 7 – 11 mm long, 2 – 2.5 mm wide; stamens 6, filaments free, dark-brown, c. 1 – 1.75 mm long, anthers elongate-lanceolate, pale creamy yellow, free, c. 5 – 7 mm long, pollen yellow, numerous, pistillodes absent. Pistillate flowers with calyx of 3 minute united sepals, c. 3 – 4 mm long, 3 – 3.5 mm wide; corolla with 3 petals, c. 6 – 8 mm long, 3 – 6 mm wide; staminodes 6, uniform, c. 1 mm long; gynoecium c. 4 mm long, 1 – 1.5 mm wide; stigma with 3 elongate lobes, c. 1 – 2 mm long, brighter coloured. Fruits globose, bilobed or trilobed, reddish orange when mature (Lauterbach 970), c. 3.3 – 5 cm diam., pale brownish green when young, orange or brownish orange when mature. Embryo placed below middle line of seed. Eophyll bifid.

distribution. Orania lauterbachiana is the most widespread species in New Guinea. It is found throughout the mainland and on a nearby island. In Papua this species is known from Sorong, Etna Bay-Fakfak and surrounding the Taritatu (formerly Idenburg) river. In Papua New Guinea it is known throughout the country from Western Province to the most eastern part (Milne Bay Province) including Normanby Island. So far, it has never been collected from other nearby islands. Map 8.

Map 8

The distribution of Orania lauterbachiana (■), O. micrantha (♦), O. clemensiae (▲) (now synonym of O. micrantha), and O. brassii (●) (now synonym of O. lauterbachiana).

specimens examined. indonesia. Papua Barat. Sorong: Ayawasi, Kecamatan Aifat, along the small stream, NW of Ayawasi, 1°09’S, 132°29’E, 12 Sept. 1995, E. F. de Vogel 9698 (BO!, K!, L!). Papua Barat. Fakfak: Teluk Etna (Etna Bay), km 45 road PT Kaltim Hutama, 2 Feb. 2001, C. Heatubun CH 336 (K!, MAN); Teluk Etna (Etna Bay), km 45 PT Kaltim Hutama logging road, Yamur, 2 Feb. 2001, R. A. Maturbongs RAM 671 (AAU, BO!, K!, LAE, MAN). Papua. Jayawijaya: Idenburg R., L. J. Brass 12966 (BO!); 4 km SW of Bernhard Camp, Idenburg R., March 1939, L. J. Brass 13375 (BO!, L!). PAPUA NEW GUINEA. North East, Gogol Mittellauf, 17 Nov. 1890, Lauterbach 970 (B holotype, FI!). Western: Fly R., 528 Mile Camp, alt. 80 m, May 1936, L. J. Brass 6610 (L!); 2 miles N of Kiunga, Upper Fly R., 11 Sept. 1967, Pullen 7303 (K!). Southern Highlands: Erave Distr., road to SE Gobe oil rig, 80 km NW of Kikori, 6°50'9.6''S 143°48'52.1''E, 26 Nov. 2000, W. J. Baker 1114 (AAU, BO!, K!, L, LAE, NY). Gulf: Kikori Distr., Victory junction (confluence of Siribi & Kuru Rs), 34 km N of Kikori, 7°7'25.9''S 144°19'30.2''E, 20 Nov. 2000, W. J. Baker 1099 (AAU, BO!, K!, LAE, NY); TFI logging concession near Morere village, 28 Km NE of Kikori, 7°15'15.1''S 144°26'20.3''E, W. J. Baker 1107 (AAU, BRI, BO!, K!, LAE, NY, L). Madang: Lower Ramu, Atitau area, 9 Oct. 1958, R. Pullen 1192 (L!). Morobe: Boridi, Mokian, 12 Nov. 1935, Carr 14884 (K!); Lae, 100 ft. alt, 3 Dec. 1958, Henty NGF 10576 (K!); Bulili ridge, near Lababia guesthouse, 7°5'S 147°5'E, 6 April 2000, A. Kjær 515 (AAU, BRI, CAN, K!). Central: Mafulu, 1933 New Guinea Expedition of the American Museum of Natural History, 12 Nov. 1933, L. J. Brass 5489 (B, BO!, NY! lectotype). Milney Bay: Normanby Island, Mt Pabinama, 10 May 1956, L. J. Brass 25820 (K!); Normanby Island, Waikaiuna, 17 May 1956, L. J. Brass 25910 (K!); Normanby Island, Mt Solomonai, W of Esa’ala, 700 m alt., 26 Nov. 1976, J. R. Croft LAE 68947 (L!).

habitat. In ridge forest from lowlands up to highland tropical forest from 10 – 820 m altitude.

conservation status. Least concern (LC). Widespread throughout the island of New Guinea and often occurring in large stands.

etymology. After C. A. G. Lauterbach, German naturalist and plant collector.

vernacular names. bananak (Amele-Madang), kolu (Mokian), kunakwan (Jal.), omoo (Yamur).

uses. The trunk is used as floorboards for houses (Burret 1935).

notes. The holotype of Orania lauterbachiana (Lauterbach 970) was once identified as O. macropetala and cited in the protologue of that species by Lauterbach & Schumann (1901) with further notes on its slight differences with the syntypes of O. macropetala, Lauterbach 2001 and 2835. In the protologue of O. lauterbachiana Beccari (1915) had already described in detail the differences between O. lauterbachiana and O. macropetala regarding the floral dimensions, but there was no mention of the number of stamens and staminodes. Neither did Lauterbach & Schumann mention this in the protologue of O. macropetala. It was Essig (1980) who first described the number of stamens as one character that distinguishes O. lauterbachiana from O. macropetala. Amazingly, although Beccari did not mention the number of stamens, his observation was correct, that Lauterbach 970 represented a different taxon. Unfortunately the holotype of O. lauterbachiana was probably destroyed during World War II. Only one isotype survived, the one kept in FI — just three photographs of rachillae and a bunch of dried staminate and pistillate flowers. Essig (1980) cited the presence of photographs of fragments of the type — presumed to be of B — at BH, but these have not been found. Essig also mentioned that O. lauterbachiana has a fruit diam. varying from 3.2 – 5.5 cm; however, the protologue of O. lauterbachiana does not say anything about the fruit diam. (see Beccari 1915). Apparently the information regarding the fruit diam. was measured from other specimens, but not the type. Burret (1936) described O. brassii as a distinct species; however, Burret (1936) never compared his O. brassii with O. lauterbachiana in his protologue although there was a great possibility that he had seen the type of O. lauterbachiana — or even read the protologue itself — sent by Beccari to Berlin several years before. Instead Burret only compared O. brassii with the glabrous O. disticha and stated the presence of red-brown tomentum in the peduncle and rachillae as the distinctive characters. The species is now subsumed, not based on the comparison with O. disticha, but with a more appropriate one, O. lauterbachiana. Essig also mentioned without giving an exact reference that Burret himself had annotated the specimen at NY, so based on this the specimen was then selected as the lectotype. This statement is doubtful for Burret never wrote about this in the protologue.

18. Orania micrantha Becc. (Beccari 1915: 36). Type: Papua New Guinea, North East, in the forest of Kani Mountain, alt. 800 m, 18 May 1908, Schlechter 17739 (holotype B†).

Orania clemensiae Burret (1940: 8). Type: Papua New Guinea, Morobe, Boana, 4000 feet (c. 1220 m) alt., 7 May 1940, Clemens 8713 (holotype B†), synon. nov.

Large palm. Trunk c. 17 m tall, c. 25 – 30 cm diam. (dbh). Leaves spirally arranged, c. 262 m long; part of rachis c. 1 – 1.3 cm diam., with dense red-brown tomentum; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, c. 30 on each side of rachis, leaflets c. 5 – 8 cm distant, c. 80 – 100 cm long, 4.7 – 6.5 cm wide, adaxial surface glabrous, red-brown tomentum on midrib, midrib thick, other ribs slender, abaxial surface densely covered with white indumentum, thin red-brown tomentum on margin, midrib thick, other ribs slender. Inflorescence spreading, branching to 2 orders, c. 100 – 270 cm long; peduncle c. 45 – 130 cm long, with dense red-brown tomentum; rachis c. 55 – 140 cm long; first order branch c. 40 – 70 cm long, rachillae bract minute or with 5 mm long, rachillae slender, c. 37 – 60 cm long, bearing 150 – 200 flower clusters, otherwise 96 – 100 (Clemens 11363), bearing regularly arranged triads in the proximal $\frac{4}{5}$ & paired staminate flowers in the distal $\frac{1}{5}$, otherwise in the proximal $\frac{1}{5}$ (Clemens 11363), the basal c. 3 – 4 cm devoid of flowers, triads c. 1 – 2 cm distant, rachilla glabrous or with thin red-brown tomentum. Staminate flowers creamy white, rarely with dark green coloured (LAE 74064); with calyx of 3 united minute sepals, c. 2 mm long; corolla with 3 free petals, c. 7 – 10 mm long, 2 – 5 mm wide; stamens 6, filaments free, dark-brown, c. 1.7 – 3.5 mm long, with one or two being longer, c. 6 – 7.5 mm long, anther elongate-lanceolate, pale creamy yellow, free, c. 4 – 5 mm long; pistillodes absent. Pistillate flowers creamy white, rarely with dark green coloured (LAE 74064); with calyx of 3 united sepals, minute or with 1.5 mm long; corolla with 3 free petals, c. 4 – 5 mm long, 2.5 – 4 mm wide; staminodes 6, uniform, c. 0.6 – 1.1 mm long; gynoecium dark-brown coloured, c. 3 – 5 mm long, 2 mm wide, stigma with 3 elongate lobes. Fruits globose or bilobed, young fruit with c. 2.8 cm diam., mature fruit unknown. Embryo placed below middle line of seed (see Essig 1980). Eophyll bifid.

distribution. This species is known from the northeast (the then Kaiser Wilhelmsland) through West Sepik to Morobe provinces in Papua New Guinea. Map 8.

specimens examined. papua new guinea. North East: Kaiser Wilhelmsland, Kani Mt, 26 April 1908, Schlechter 17610 (B, FI!); 18 May 1908, Schlechter 17735 (B, FI!). West Sepik: Telefomin, Carpentaria Exploration Co., Frieda R. site, Antap Mt, helicopter pad, 1380 m alt., 29 April 1978, Essig & Young LAE 74064 (BH, L!, LAE). Morobe: Boana, 7 May 1940, Clemens 11363 (K!).

habitat. Highland tropical forest with an altitude of 800 – 1380 m.

conservation status. Data deficient (DD). The palm is known only from the type locality and two other collections. We have no recent information on the status of forests in these localities.

etymology. With small flowers.

vernacular names. Unknown.

uses. Unknown.

notes. Orania micrantha is probably the most elusive species of Orania. In the protologue of this species Beccari (1915) selected the specimen in Berlin as the holotype, this was probably destroyed during the Second World War. Essig (1980) in his monograph mentioned that he had seen photographs of the fragments of the type in FI and BH. Unfortunately we did not see any photographs of the types in FI, instead we did see two photographs and fragments of rachilla identified by Beccari himself as O. micrantha but with different collection numbers — definitely not of the type which is Schlechter 17739. One photograph was apparently of Schlechter 17735, the other one was unlabelled and there is no information about the collection number written on it except that it was attached with the specimen of Schlechter 17610. It seems to have been removed from the herbarium specimen. We presumed that this photograph is of that particular collection, not of the type. On the other hand, the fragment of rachilla was clearly labelled each for Schlechter 17735 and 17610. Looking at those two photographs and fragments, in general we found no significant difference in appearance. We could not find the type specimen in FI, nor even the fragment of it as was previously stated by Essig (1980). Efforts were made to borrow the photographs — or at least copies of them — from BH as a proof that they do exist somewhere outside B and FI, unfortunately without success. Therefore, all the information needed to determine the status of this species relies only on those two non-type specimens available in FI. The new specimen collected by Essig (Essig & Young LAE 74064) which he believed may represent a new species or just a subspecies of O. lauterbachiana, based on this study turns to be a member of this species. The type of the then O. clemensiae was most likely destroyed during the Second World War and there is no information on the existence of a duplicate kept in other herbaria. Thus it is only known through its protologue and the single collection under a different collection number made by the same collector and year from approximately the same location as the type, available at Kew. The results of this study indicate that there is no single distinctive morphological character, which can be used to separate O. clemensiae from O. micrantha as two distinct species, thus O. micrantha is synonymised. This is not in accordance with Burret (1940). In his monograph Essig (1980) regarded this species as a synonym of O. lauterbachiana based on observations made from the new specimen that he collected from the type locality, without mature flowers and fruits (Essig & Young LAE 74012).

The subgroup ‘Macropetala complex’

Three species belong to this complex, Orania bakeri, O. grandiflora and O. macropetala, based on the similarity in the size of their pistillate flowers. Differences are shown in Table 16.

Table 16 Morphological features distinguishing Orania bakeri, O. grandiflora, and O. macropetala.

19. Orania bakeri A. P. Keim & J. Dransf. sp. nov.

Palma grandis, O. macropetalae simulans sed 40 ad 55 fasciculis florum per rachillam, floribus staminatis petalis c. 10 mm longis, staminibus 6, floribus pistillatis petalis c. 15 mm longis, staminodiis 6 fructibus atrobrunneis, c. 4 cm diametro differt. Typus: Papua New Guinea, Madang, Goinbang, near Bosmun, Bogia, 18 Jan. 1996, W. J. Baker 581 (holotypus K!; isotypus LAE).

http://www.ipni.org/urn:lsid:ipni.org:names:77118914-1

Large palm. Trunk c. 15 m tall. Leaves 8 in the crown, spirally arranged, c. 4 – 5 m long; leaf-sheath massive, c. 30 cm long, 14 cm wide, adaxial surface glabrous, abaxial surface slightly covered with red-brown tomentum, margins massively disintegrating into fibres, fibres straight, c. 13.5 – 14 cm long each; petiole 1.1 m long, covered with red-brown tomentum; rachis c. 2.6 – 3.6 m long, in middle part c. 2.5 cm diam.; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, c. 40 – 45 in each side of rachis, in middle part of rachis distance between 2 leaflets c. 5.5 – 7.5 cm, c. 1.32 m long, 6 – 7 cm wide, adaxial surface glabrous, midrib thick, other ribs slender, abaxial surface slightly covered with white indumentum, midrib robust, other ribs slender but thicker than adaxial. Inflorescence spreading, branching to 2 orders; prophyll small, persistent, disintegrating into fibres; peduncle c. 40 cm long, with thin red-brown tomentum; peduncular bract 1, woody, c. 1.15 – 1.4 m long, including 32 cm acumen, 9 cm wide at near base, splitting in the middle, disintegrating into fibres when old, adaxially glabrous, abaxially with dense red-brown tomentum; first order branch c. 48 – 55 cm long, rachillae bract c. 8 – 9 mm long, rachillae robust, not conspicuously zigzag, c. 40.5 – 55 cm long, bearing 40 – 55 flower clusters, bearing triads on about proximal $\frac{1}{2}$ and paired staminate flowers in the distal $\frac{1}{2}$, the basal c. 2 cm devoid of flowers, triads c. 2 cm distant, rachilla with greyish white indumentum. Staminate flowers with calyx of 3 united minute sepals; corolla with 3 free petals, c. 10 × 3 mm; stamens 6, filaments free, dark brown, c. 1.7 – 2.2 mm long, anthers elongate-lanceolate, pale creamy-yellow, free, c. 6.7 mm long; pistillodes absent. Pistillate flowers with calyx of 3 united sepals, c. 3 mm long; corolla with 3 free petals, c. 15 × 7 mm, yellowish cream, thick; staminodes 6, c. 3.5 cm long; gynoecium dark brown, c. 7 × 6 mm; stigma with 3 elongate lobes. Fruit globose or bilobed, c. 3.8 – 3.9 cm diam., olive green when young, dark brown when mature (W. J. Baker 1999 pers. comm.), stigmatic remains sub-basal; epicarp thin, smooth; mesocarp fibrous, c. 4 – 5 mm thick; endocarp hard, red-brown coloured, c. 2 – 3 mm thick; testa hard, 1 – 2 mm thick; endosperm white, c. 2.4 cm diam., with hollow inside. Embryo placed below middle line of seed. Eophyll bifid. Fig. 7.

distribution. Endemic to a small area in the eastern part of New Guinea, especially in the province of Madang. Map 9.

Map 9

The distribution of Orania zonae (●), O. longistaminodia (■), O. gagavu (▲), O. glauca (♦), O. grandiflora (▼), O. bakeri (◊), and O. deflexa (►).

specimens examined. papua new guinea. Madang: Road between Bogia and Bosmun 2, Goinbang, near Bosmun 2, mouth of Ramu R. area, 18 Jan. 1996, W. J. Baker 581 (K!, holotype; LAE).

habitat. Rather swampy in lowland coastal tropical rainforest at an altitude of about 50 m above sea level.

conservation status. Critically endangered (CR B1 B2). The species occurs in considerable numbers at its only known locality but the area is extensively logged and the estimated extent of occurrence is less than 100 km². Certainly the quality of the habitat is in decline (W. J. Baker, pers. comm.).

etymology. After William J. Baker, British palm taxonomist and collector of the type specimen.

vernacular names. Unknown.

uses. Unknown.

notes. Apart from the thickness of the rachillae, number of stamens and staminodes, this species looks very similar to Orania macropetala. The distribution area of O. bakeri itself overlaps the distribution area of O. macropetala. It is just unfortunate that we have only one specimen of O. bakeri with fewer than 10 staminate flowers. Nevertheless, O. bakeri is regarded here as a distinct species from O. macropetala based ultimately on the fact that all specimens of O. macropetala observed in this current study, including the paratype (Lauterbach 2001), all possess nine to 14 stamens.

20. Orania grandiflora A. P. Keim & J. Dransf. sp. nov.

Palma grandis, O. macropetalae simulans sed rachillis rubro-brunneis tomento dense tectis, 63 – 70 fasciculius florum per rachillam, floribus staminatis petalis circa 9 ad 10 mm longis, staminibus 6; floribus pistillatis petalis c. 6 ad 8 mm longis, staminodiis 6, fructibus aurantiacis vel rubro-aurantiacis, c. 5 cm diametro differt. Typus: Indonesia, Papua, Fakfak, road down E Levee towards Kampung Kali Kopi, loc.10, Timika, 10 Feb. 1998, C. Heatubun CH 179 (holotypus K!; isotypus MAN!).

http://www.ipni.org/urn:lsid:ipni.org:names:77118915-1

Large palm. Trunk c. 25 m high. Leaves 10 in the crown, spirally arranged, c. 6 m long; leaf-sheath massive, adaxial surface glabrous, abaxial surface densely covered with red-brown tomentum and creamy white indumentum, margins disintegrating into fibres, fibres short, straight; petiole 2 m long, c. 4.5 – 5 cm diam., red-brown tomentum present; rachis c. 3 cm diam. in the middle part, with dense red-brown tomentum and creamy white indumentum; leaflets elongate-lanceolate, totally 58 – 60, regularly arranged leaflets held in one plane, the proximal 2 leaflets crowded 1.5 – 2 cm distant, otherwise c. 7 cm distant in the middle part of rachis, c. 1.5 m long, 7.5 cm wide, adaxial surface glabrous, midrib robust, with sparse red-brown tomentum, other ribs slender, glabrous, abaxial surface densely covered with white indumentum, wax evidence present, red-brown tomentum absent, midrib less robust than adaxial, other ribs thicker than adaxial. Inflorescence spreading, branching to 2 orders, robust, densely covered with red-brown tomentum; prophyll hard, persistent, disintegrating into fibres when old, c. 30 cm long, 15 cm wide near the base, adaxial and abaxial surface glabrous; peduncular bract one, woody, adaxial surface glabrous, abaxial surface with dense red-brown tomentum, splitting in the middle, heavily disintegrating into fibres, persistent but shorter than prophyll; first order branch c. 80 – 85 cm long, rachillae bract scale-like; rachillae conspicuously robust, proximally c. 1 – 1.2 cm diam., with dense red-brown tomentum, c. 54 – 63 cm long, bearing 63 – 70 flower clusters, bearing triads arranged conspicuously distichously in the proximal $\frac{1}{2}$ and paired staminate flowers in the distal $\frac{1}{2}$, the basal c. 2 cm devoid of flowers, triads c. 1.5 cm distant, rachilla with dense red-brown tomentum. Staminate flowers with calyx of 3 united minute sepals; corolla with 3 free petals, c. 9 – 10 mm long, 2.5 – 3 mm wide; stamens 6, filaments always free, dark brown, c. 1 mm long, anthers elongate-lanceolate, c. 4 mm long, pale creamy yellow, pistillode absent. Pistillate flowers with calyx of 3 united sepals, c. 2.5 – 3 mm long; corolla with 3 free petals, c. 10 mm long, 6 – 6.5 mm wide; staminodes 6, c. 2 mm long, always uniform; gynaecium, c. 5 – 6 mm long, 4 mm wide; stigma with 3 elongate lobes, c. 2.5 – 3 mm. Fruit globose or bilobed, c. 4.8 – 5 cm diam., mature fruit orange or reddish orange (W. J. Baker 1999 pers. comm.), stigmatic remains sub basal, smooth surface. Embryo placed below middle line of seed. Eophyll bifid. Fig. 8.

distribution. Endemic to Papua. Map 9.

specimen examined. indonesia. Papua. Fakfak: Road down E Levee towards Kampung Kali Kopi, loc. 104.26°82'S and 136.5°5.9'E Timika, 10 Feb. 1998, C. Heatubun CH 179 (K! holotype, MAN!).

habitat. Disturbed alluvial forest with altitude about 30 m.

conservation status. Data deficient (DD). The palm is known only from its type. In 1999 the locality was at the side of a bund next to major tailings of the P. T. Freeport Copper mine, a locality extremely vulnerable to disturbance. We do not know whether the population of the palm is still extant.

etymology. With large flowers.

vernacular names. Unknown.

uses. Not recorded.

notes. Orania grandiflora is distinguished from O. macropetala primarily by the presence of dense red-brown tomentum in the inflorescence, size of both staminate and pistillate flowers and the numbers of stamens and staminodes (Table 16). Unlike O. bakeri, the distribution of O. grandiflora does not overlap the distribution of O. macropetala. O. macropetala has never been collected from the southern part of New Guinea mainland. However, this may be caused by undercollecting of a possible widespread species. There have been limited collections of O. macropetala made from this part of mainland New Guinea. Further study with more collecting is essential.

21. Orania macropetala K. Schum. & Lauterb. in Lauterbach & Schumann (1901: 205). Type: Papua New Guinea, Bismarck Mountain, 10 Sept. 1896, Lauterbach 2835 (lectotype B†; isolectotype FI!, selected here).

Large palm. Trunk 10 – 20 m high, c. 20 cm diam. (dbh). Leaves c. 12 in the crown, spirally arranged, c. 3.9 – 5 m long; leaf-sheath 42 – 50 cm long, adaxial surface glabrous, abaxial surface with dense red-brown tomentum, margins disintegrating into fibres; petiole c. 1 m long; rachis c. 2.9 – 4 m long, c. 2.5 cm diam. in the middle part; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, c. 6.5 – 7 cm distant; c. 79 – 140 cm long, 5 – 8.5 cm wide, adaxial surface glabrous, midrib robust, other ribs slender, glabrous, abaxial surface densely covered with white indumentum, thin red-brown tomentum in margin and basal part, midrib massive, other ribs slender, glabrous. Inflorescence spreading, branching to 2 orders, robust, c. 200 cm long (J. R. Croft et al. LAE 71098); peduncle glabrous or with red-brown tomentum, sometimes with creamy white indumentum, c. 100 cm long (J. R. Croft et al. LAE 71098); first order branches c. 60 – 70 cm long, sometimes c. 10 cm, rachis bract c. 5 – 1.1 cm long, 3.5 – 1.5 cm wide, rachillae robust, conspicuously zigzag, with greyish white indumentum, c. 30 – 50 cm long, bearing 30 – 50 flower clusters, bearing triads in the proximal $\frac{4}{5}$ and paired staminate flowers in the distal $\frac{1}{5}$ part, the basal c. 2 – 4 cm devoid of flowers, triads c. 2 – 3 cm distant. Staminate flowers with calyx of 3 united sepals, c. 1.5 – 4 mm long; corolla with 3 free petals, c. 15 – 17 mm long, 4.5 – 5 mm wide; stamens 9 – 14, filaments always free, dark brown, c. 1 – 2 mm long, anthers elongate-lanceolate, pale creamy yellow, free, c. 5 – 7 mm long, pistillode absent. Pistillate flowers with calyx of 3 united sepals, c. 1 – 2 cm long; corolla with 3 free petals, c. 9 – 13 mm long, 5 – 6 mm wide; staminodes 6 or 10, c. 1.5 – 3 mm long, uniform; gynoecium c. 6 × 5 mm, stigma with 3 elongate lobes, brighter coloured, c. 2 mm long. Fruits globose, c. 6 cm diam.; epicarp thin, smooth; mesocarp fibrous, c. 3 – 4 mm thick; endocarp red-brown, hard; endosperm white, c. 5 cm diam., 1.4 cm thick, with hollow inside, c. 2 cm diam. Embryo placed below middle line of seed. Eophyll bifid.

distribution. New Guinea in both the Indonesian Province of Papua Barat and Papua New Guinea. In Papua Barat it is only known from a limited area around Manokwari. In Papua New Guinea it is found in the northern part especially between Madang and Morobe Provinces. Map 6.

specimens examined. indonesia. Papua Barat. Manokwari: Waren, 60 miles S of Manokwari, 2 March 1940, Kanehira & Hatusima 12967 (BO!). PAPUA NEW GUINEA. Ebene: Astrolobe, May 1896, Lauterbach 2001 (FI! paratype); Bismarck Mountain, 10 Sept. 1896, Lauterbach 2835 (FI! isolectotype). Northern/Kaiser Wilhelmsland: Woildtum to Djamu, 20 April 1908, Schlechter ‘Djamu’ (L!); Djamu, 20 April 1908, Schlechter 17580 (FI!). Morobe: Kajabit, Aug. – Dec. 1939, Clemens 10894 (K!); Finschhafen, Banario Mountain, near Nanduo village, 28 km NW of Finschhafen, 6°25'55.5''S 147°40'3.2''E, 6 Dec. 2000, R. Banka 2014 (BO!, K!, LAE). Madang: Bongu, 1978, J. R. Croft et al. LAE 71098 (L!); Mountain 2 km S of Usino, 5 April 1978, Essig & Young LAE 74001 (BH, L!, LAE).

habitat. From lowland to highland tropical rainforest within altitude range 100 – 900 m above sea level.

conservation status. Near threatened (NT). Although widespread in New Guinea, most of the collections were made at least 40 years ago with only one collection made recently.

etymology. Big petals.

vernacular names. gamain (Usino-Madang).

uses. Unknown.

notes. Essig (1980) noted that the number of staminodes in Orania macropetala is three or four. We cannot account for this description because the protologue (Lauterbach & Schumann 1901) does not mention it — not even the nine to 14 stamens. Both syntypes in FI (Lauterbach 2001 and 2835) are without flowers. Actually they are available only as parts of leaf rachis, rachillae and part of mature fruit — fortunately Lauterbach 2835 is accompanied by two clear photographs — but nothing could be done to observe the staminodes. The observation of flowers then was based on the three non-type complete specimens, which match the protologue. Two specimens show six staminodes, the other ten. Beccari (1915) selected a specimen of O. macropetala with both staminate and pistillate flowers Schlechter’s ‘Djamer’ (Djamu?) no. 17980 kept in B. Unfortunately this specimen was most likely destroyed during the Second World War. In FI there is one specimen of O. macropetala collected by Schlechter from near a forest called ‘Djamu’ in the then German New Guinea, Schlechter 17580, with an annotation that the specimen is a copy or duplicate sent to FI by B. Until we can trace Schlechter’s original ‘Djamer’ 17980, we presume that this specimen is actually the one which Beccari mentioned. The number 17980 might be just a misprint of 17580. Another ‘Djamu’ specimen is kept in L and without collection number although it was also collected by Schlechter on the same day as Schlechter 17580. The FI one has both ten stamens and staminodes. The L one has 11 stamens and just six staminodes. Could the L specimen be the duplicate of Schlechter ‘Djamer’ 17980? According to the protologue, the mature fruits are 7 – 7.5 cm diam. — presumably there was fresh material (from dried materials we measured only 6 cm). If they are, O. macropetala could be one of the four New Guinean taxa, which have large fruits — the other three are O. longistaminodia, O. palindan and O. regalis. The selection of Lauterbach 2835 as lectotype by Essig is accepted for this specimen at FI; it has the more significantly representative appearance than the other one — Lauterbach 2001. O. macropetala is one of three Malesian species with more than six stamens; the others are O. longistaminodia and O. zonae. However, these three species are different in several characters (Table 17). O. macropetala in some way resembles O. lauterbachiana, especially regarding the robust rachillae and the size and shape of staminate flowers. However, they differ in several characters (Table 18).

Table 17 Morphological features distinguishing Orania longistaminodia, O. macropetala and O. zonae.

Table 18 Morphological differences between Orania lauterbachiana and O. macropetala.

The subgroup of species with ‘Sylvicola type’ inflorescences

Orania glauca, O. regalis and O. sylvicola are grouped together here on the basis of similarity in their inflorescences. For details on ‘Sylvicola type’ inflorescence, see the section ‘Inflorescences’ see p. 132. However, despite the similarity, the three species are obviously distinct (Table 19).

Table 19 Morphological features distinguishing Orania glauca, O. regalis, and O. sylvicola.

22. Orania glauca Essig (1980: 229). Type: Papua New Guinea, West Sepik, near the city of Amanab, 24 Nov. 1971, Essig LAE 55089 (holotype BH!; isotype LAE).

Robust palm. Trunk c. 30 cm diam. (dbh). Leaves c. 8 – 10 in the crown, arranged spirally, wax conspicuously present, c. 4.6 m long; leaf-sheath and petiole 1.4 m long, margins disintegrating into fibres, adaxial surface glabrous, abaxial surface slightly covered with red-brown tomentum; rachis 3.2 m long, c. 3 cm diam. in the middle, with slightly red-brown tomentum; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, 60 on either side of the rachis, c. 80 – 84 cm long, 6.8 – 7 cm wide, adaxial surface dark green, glabrous, with conspicuous wax on terminal leaflets, midrib robust, with wax, other ribs less thick, glabrous, abaxial surface slightly covered with white indumentum and wax, midrib robust, secondary ribs thick, other ribs less thick. Inflorescence spreading, branching to 3 orders, glabrous, wax conspicuous, with lightly glaucous appearance, c. 260 cm long; peduncle c. 80 cm long; peduncular bract one, persistent, woody, wax conspicuous, indumentum unclear, c. 3 m long, splitting in the middle; rachis 180 cm long, rachillae branching at convergent angle, conspicuously slender, not conspicuously zigzagging, c. 53 first order branches, c. 80 cm long, second order branches c. 15 cm long, rachillae short, c. 13.5 – 15 cm long, triads arranged in the proximal c. $\frac{4}{5}$ part, the basal c. 3 cm devoid of flowers, triads c. 1.5 cm distant, flower clusters 12 – 15 in a rachilla, proximally c. $\frac{1}{2}$ – $\frac{4}{5}$ contains pistillate flowers. Flowers unknown. Fruits globose, c. 4.0 – 4.8 cm diam., bi- and tri-lobed form present, stigmatic remains sub-basal, dull green when young, reddish yellow to pale orange when mature. Embryo placed below middle line of seed. Eophyll bifid.

distribution. Papua New Guinea: endemic to the type location. Map 9.

specimen examined. papua new guinea. West Sepik: Near the city of Amanab, 3°40'S, 141°15'E, 24 Nov. 1971, Essig LAE 55089 (BH! holotype, LAE).

habitat. Disturbed red soil forest.

conservation status. Data deficient (DD). The palm is known only from its type and we have no recent information on the status of forests at the type locality.

etymology. Glaucous — appearance of the inflorescence.

vernacular names. Unknown.

uses. Unknown.

notes. Essig (1980) stated that this species is one of the largest members of the genus. His statement is supported by this present study. Within the three species in this subgroup, Orania glauca has one unique character; the possession of an inflorescence branched to three orders. The other two Malesian species that possess this character are O. deflexa and O. gagavu; differences are shown in Table 20.

Table 20 Comparison between Orania deflexa, O. gagavu and O. glauca.

23. Orania regalis Zipp. ex Blume (1836: 116, t. 119 & 122). Type: Arausiaca excelsa Blume.

Orania regalis Zipp. (in Blume 1829: 297). nom. nud.

Arausiaca excelsa Blume (1836: 8, t.119 & 122).

Orania aruensis Becc. (Beccari 1877: 76). Type: Indonesia, Moluccas, Aru Archipelago, Wokam Island, Jabulenga, Beccari s.n. ‘Giabu Lengan’ (holotype FI!; isotypes BO!, K!, L!), synon. nov.

Large palm. Trunk up to 11 m high, c. 20 – 25 cm diam., internodes 15 – 20 cm, glaborus, greyish green. Leaves 8 – 10 in the crown, spirally arranged, c. 2.5 – 4.2 m long; leaf-sheath c. 60 cm long, 18 cm wide, margins disintegrating into fibres, fibres curly, c. 10 cm long; petiole about 4 – 4.5 cm diam. in the proximal, with dense red-brown tomentum; rachis c. 2 cm diam., leaflets elongate-lanceolate, regularly arranged, 50 – 60 in one side of rachis, leaflets c. 5.0 – 6.0 cm distant, c. 1 – 1.6 m long, 6.5 – 7.2 cm wide, adaxial surface dark green, glabrous, with red brown tomentum on the midrib, midrib robust, other ribs slender, abaxial surface with dense white indumentum, red-brown tomentum on the margin, midrib robust, other ribs rather slender, more conspicuous than adaxial ones. Inflorescence congested, branching to 2 orders, c. 1.20 – 1.25 m long; prophyll persistent, c. 1 – 1.2 m long, heavily disintegrated into fibres when old; peduncle 60 – 62.5 cm long, glabrous, yellowish green; peduncular bract one, woody, c. 83 – 150 cm long, 10.5 – 11 cm wide, adaxial surface glabrous, abaxial surface with red-brown tomentum, splitting in the middle, proximally disintegrating into fibres; rachis c. 60 – 62.5 cm long, glabrous, yellowish green, rachis bract c. 2.5 – 3 mm long, 7 – 8 mm wide; first order branches c. 19 – 20 cm long, rachillae c. 12 – 20 cm long, slender, conspicuously zigzag, branching at convergent angle, bearing 30 – 60 flower clusters, bearing triads spirally arranged in the proximal $\frac{2}{5}$, the basal c. 1.2 – 1.6 cm devoid of flowers, triads c. 1 – 1.5 cm distant, rachilla glabrous. Staminate flowers with calyx of 3 united minute sepals; corolla with 3 free petals, c. 4 – 7 mm long, 1 – 2 mm wide; stamens 3 – 5, commonly 3 (–rarely more), filaments free or united, dark brown, c. 1 mm long, anthers pale creamy yellow, c. 2.5 – 3 mm long, free or united, uniform or distinct, pistillodes absent. Pistillate flowers with calyx of 3 united sepals, c. 1 – 1.5 mm long; corolla with 3 free petals, c. 5 – 7 mm long, 3 – 4 mm wide; staminodes 3, c. 0.4 – 0.5 mm long; gynoecium brown, c. 2 mm long; stigma with 3 elongate lobes, c. 1 mm long. Fruit globose or bilobed, dull green when young, bright/reddish orange when mature, c. 4.5 – 8.0 cm diam.; epicarp smooth, thin; mesocarp fibrous, c. 4 mm thick; endocarp thinner than mesocarp but thicker than epicarp, woody, brown, c. 1 mm thick; endosperm c. 3.5 – 3.6 cm diam., with a hollow, c. 1 cm diam. Embryo placed below middle line of seed. Eophyll bifid.

distribution. Moluccas and New Guinea. In the Moluccas it is found near the modern town of Jabulenga on the island of Wokam, in the northern part of the Aru Archipelago. In New Guinea it is found so far only in Papua in both Northern and Southern parts of the ‘neck’. In the northern part it is found around Wandammen National Park — about 80 miles northwest of Triton Bay (Baker et al. 2000) and Nabire — approximately 90 miles west of Triton Bay (Heatubun 1999 pers. comm.). In the southern part it has not been seen since Zippelius’s original collection in 1827. Zippelius mentions in his notes its presence in the mountain of Lamentsjerie close to a town called Lobo (present day Dobo?) not very far from Triton Bay and the district capital city Kaimana. Map 7.

specimens examined. indonesia. Moluccas. Aru Archipelago: ‘Giabu-lengan’ (Jabulenga), ‘Vokan’ (Wokam) Island, May 1873, Beccari s.n. (BO!, FI!, K!, L!). Papua Barat. Sorong: Misoöl, Limalas Village, 19 Jan. 2002, R. A. Maturbongs RAM 693 (BO!, K!, LAE, MAN); Motlol, 22 Jan. 2002, C. Heatubun et al. CH 356 (K!, MAN). Manokwari: Senderawoi (Wosimi R.), 25 Feb. 2000, C. Heatubun CH 317 (AAU, BO!, K!, MAN); Wandammen Peninsula, Wasior distr., near Dotir village, 11 km N of Wasior, near the confluence of the Mawoi and Yois Rs., 2000, W. J. Baker 1045 (BO!, K!, MAN). Papua. Paniai: Nabire, 1998, C. Heatubun CH 297 (MAN!); Samabusa, 10 Feb. 2001, R.A. Maturbongs RAM 675 (BO!, K!, MAN). CULTIVATED. Java: Bogor Botanic Garden, bed II.F.5, 6 May 1902, J. C. Schoute s.n. (BO!, L!); Anonymous s.n. Communic. ex. Herb. Hort. Bot. Bog. no. 12 (BO!, L!); bed XII.E.21a, 16 Jan. 1999, A. Keim AK 32 (BH!, K!); bed XII.E.21, 16 Jan. 1999, A. Keim AK 33 (K!, MAN!); bed V.H.63, 17 June 1999, A. Keim AK 47 (BO!, K!); bed V.A.53, Anonymous s.n. (L!); bed XII.E.21, Anonymous s.n. (FI!). Singapore: Singapore Botanic Garden, Lawn o, 8 Oct. 1929, Nur s.n. (K!, SING); Lawn o, 10 Jan. 1933, Kiah SF 26189 (BO!, K!, SING). Sri Lanka: Peradineya Botanic Garden, 23 July 1886, Rutherford et al. R-146 (K!).

habitat. Lowland to highland tropical rainforest.

conservation status. Vulnerable (VU D2). Although Orania regalis has a wide extent of occurrence, only four unique localities have been recorded and the lowland forests where the palm occurs continue to be threatened by clearance for agriculture and logging.

etymology. Royal.

vernacular names. Unknown.

uses. Unknown.

notes. The presence of Orania regalis on Wokam Island, an island within the Aru Archipelago, is not surprising as the Aru Archipelago was connected to mainland New Guinea until some 10,000 years ago (Van Balgooy 1996). Concerning the nature of stamens, the protologue of both O. regalis and then O. aruensis clearly mentioned that the number is three with the anthers always free and so are the filaments. However, observations on a herbarium specimen from Bogor Botanic Garden kept in FI collected from bed no.XIIE.21, without date (Anonymous s.n.) reveal a different situation. In this specimen the number of stamens varies from three to five, but never more than five. Some of the anthers are united in pairs giving the appearance of three stamens. The filaments are always united. The herbarium specimens recently collected from living collections in Bogor Botanic Garden planted in bed XIIE.21 and 21a (AK32 and 33) have those united anthers and filaments. The same is also the case in the herbarium specimen taken from another living collection, which was still labelled as O. aruensis, planted in bed VH.63 (A. Keim AK 47). Observations made on herbarium specimens kept in the botanic garden labelled as collected from locations XII.E.21 and 21a by various collectors from different years of collecting show the united anthers and filaments. At first it was thought that those exceptions only occur in Bogor Botanic Garden as they were not found in herbarium specimens collected from other botanic gardens. However, the same phenomenon is also found in the most recently collected material from Wandammen Peninsula (W. Baker 1045). The specimen (C. Hetaubun CH 297) collected from Nabire, not very far from the Wandammen Peninsula has three stamens. Despite the difference in stamen number, the rest of the morphology exactly matches with the protologue of O. regalis. The presence of O. regalis in Bogor Botanic Garden was first recorded by Blume (1823) in the first edition of the list of plant collections in the botanic garden, unfortunately without the exact location (bed) where it was planted. Subsequent lists (Hasskarl 1844); Teijsmann & Binnendijk 1866) also record O. regalis but without location. The exact location of the collections of Orania in the botanic garden was mentioned in the fourth edition of the list (Anonymous 1892) as planted in beds II.F, II.J, X.D, XII.E and XIII.A. Unfortunately, neither O. regalis nor O. aruensis was mentioned. Boldingh (1914) wrote the fifth edition of the list and for the first time the exact bed of O. regalis (as O. aruensis) was mentioned, bed XII.E.21. An individual of O. regalis is still found in this bed. The detailed location in the garden where O. regalis was planted was mentioned in the sixth edition of the list (Dakkus 1930). In this edition O. regalis (as O. aruensis) was mentioned to be planted in II.F.5 – 5a. The other locations were in X.D 35 – 35a, XII.E.21 – 21a. O. regalis can still be found in these beds with the addition of V.H.63. We regard O. aruensis as a synonym of O. regalis, in contrast to previous workers (Table 1). In the protologue of O. aruensis, Beccari (1877) himself mentioned the similarity and suggested an affinity, between the two taxa. Beccari stated that they differ mainly in their leaflets. In O. aruensis the apex of the leaflet is acuminate, but not in O. regalis. The middle nerve in O. aruensis continues to the apex of leaflet, but not in O. regalis. The rest of the description matches Orania regalis, including the obviously congested inflorescence and three stamens. The holotype of O. aruensis shows no differences from the description in the protologue of O. regalis, thus O. aruensis is no longer recognised here as distinct from O. regalis. O. regalis shares the presence of united anthers and filaments with O. zonae; however, these two species are straightforwardly distinct (Table 21).

Table 21 Morphological differences between Orania regalis and O. zonae.

24. Orania sylvicola (Griff.) H. E. Moore (1962: 44). Type: Macrocladus sylvicola Griff.

Macrocladus sylvicola Griff. (Griffith 1845: 490). Type: Malaysia, West Malaysia, Malay Peninsula, 1850, Griffith collection HEJC (holotype K!)

Orania macrocladus Mart. (Martius 1849: 186, t.177, ed. 2, nom. illeg.). Type: Macrocladus sylvicola Griff.

Large palm. Trunk c. 15 m high, c. 15 cm diam. (dbh), internodes c. 8 cm. Leaves c. 15 in the crown, spirally arranged, densely covered with red-brown tomentum, c. 4 – 5 m long; leaf-sheath c. 80 cm, 8 – 10 cm wide, adaxial surface glabrous, abaxial surface with dense red-brown tomentum, margins disintegrating into fibres, straight, c. 5 cm long; petiole c. 1 m long; rachis 2.2 – 3.2 m long, c. 3 cm diam. in the middle; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, c. 100 in total number, leaflets c. 5.5 – 6 cm distant, c. 1.05 – 1.5 m long, 6.8 – 7 cm wide, adaxial surface dark green, glabrous, with red-brown tomentum on the midrib, midrib robust, other less thick; abaxial surface densely covered with white indumentum, red-brown tomentum on the margin, midrib robust, other ribs less thick. Inflorescence spreading, branching to 2 orders, massive, rather glabrous or with red-brown tomentum, c. 1.5 – 2 m long; prophyll persistent, disintegrating into fibres, c. 45 × 12 cm; peduncle c. 75 – 100 cm long, glabrous or with white indumentum; peduncular bract one, woody, persistent, c. 2 – 2.5 m long, 10 – 11 cm wide near the base; rachis c. 75 – 100 cm long; first order branches c. 41 – 55 cm long, rachillae bract c. 3 – 4 mm long; rachillae conspicuously slender, branching at convergent angle, conspicuously straight, c. 30 – 42 cm long, bearing 100 – 154 flower clusters, bearing triads arranged in proximal c. $\frac{2}{3}$ up to 2.5 cm from distal, triads c. 7 – 15 mm distant, the basal c. 1 – 1.7 cm devoid of flowers, sometimes pistillate flowers found in the first order branch. Staminate flowers with calyx of 3 united minute sepals; corolla with 3 free petals, c. 5 – 6 mm long, 2 – 2.5 mm wide; stamens 6, filaments free, dark-brown, c. 0.5 – 0.75 mm long, anthers elongate-lanceolate, pale creamy yellow, always free, c. 2.5 – 3 mm long; pistillode absent. Pistillate flowers with calyx of 3 united sepals, c. 0.5 – 0.7 mm long; corolla with 3 free petals, c. 3.5 – 4 mm long, 3 – 4 mm wide; staminodes 6, unequal, 2 being different, larger with hooked tip, c. 10 mm long, otherwise c. 1.2 – 2 mm long; gynoecium dark-brown, c. 3.5 – 4 mm long; stigma with 3 elongate lobes. Fruits globose or bilobed, c. 4.5 – 5 cm diam., dull green when young, yellowish green when mature; epicarp smooth, thin; mesocarp fibrous, 1 – 1.5 mm thick; endocarp thinner, hard, red-brown; endosperm white, c. 3 – 4 cm diam., c. 2.5 cm thick, with a hollow inside, c. 1 – 1.5 cm wide. Embryo placed below middle line of seed. Eophyll bifid.

distribution. Widespread in almost every part of Western Malesia and slightly beyond the boundary with Indochina floral region (restricted only in southern part of Thailand). Area of distribution covers Malay Peninsula, Singapore, Sumatra, West Java, Anambas Islands, Karimata Islands Group, West Kalimantan and Sarawak. The species so far has never been recorded in the Philippines, Sulawesi and Sabah. Map 10.

Map 10

The distribution of Orania sylvicola (●)

specimens examined. thailand. Kao Salaw Kwabi, 26 March 1927, A. F. G. Kerr 12440 (K!); Klaung Ton Satul, 1927, A. F. G. Kerr 14460 (K!); Tamben Kao Panom-Ikrabi, 31 March 1930, A. F. G. Kerr 18802 (K!); Thalae Ban, 20 Km NE of Satun, 5 Nov. 1990, Barfod et al. 41133 (K!). MALAYSIA. West Malaysia. Malay Peninsula: 1850, Griffith collection HEJC 1850 (K! holotype). Pahang: Sedagong, Pulau Tioman, 24 March 1929, Nur 21747 (BO!). Perak: Perak, Scortechini 247b (FI!). East Malaysia. Sarawak: Kampong Piching, First Division, 16 May 1981, J. Dransfield JD 6061 (K!); Kampung Moglos, 18 March 1989, Laitun anak Daud KP $\frac{{55}}{{11}}$ (K!). SINGAPORE. Chan Cha Kai, 1891, Ridley 3144 (FI!, K!); Pulau Ubin, 1891, Ridley 3146 (FI!, K!). INDONESIA. North Sumatra: Pulo Liman, Padang Sidempuan, 29 Aug. – 3 Sept. 1933, Rahmat Sitoroes 5330 (K!, L!); Padang Lawas, 21 – 27 Sept. 1933, Rahmat Sitoroes 5568 (L!); Pargambiran, Asahan, 18 – 30 Oct. 1933, Rahmat Sibuea 5826 (K!); Sibolangit, Boven Bila, 14 April 1923, J. A. Lörzing 9679 (BO!, L!); Serdang, Gallia Estate above Bangun Purba, 19 Nov. 1928, J. A. Lörzing & van Vreeden 14605 (BO!). North West Sumatra: Anonymous s.n. (FI!). Jambi: Batang Ule, Muara Bungo, July 1992, Valerie Trichon 132 (K!). South Sumatra: Palembang, 27 Feb. 1892, Burmann van Vreeden 160 (BO!). West Java: Depok, near Jakarta, 24 Aug. 1898, S. H. Koorders 31040b (BO!); 16 April 1903, S. H. Koorders 40188b (BO!); 18 April 1903, S. H. Koorders 40190b (BO!, FI!); 14 March 1929, Van Steenis 2836 (BO!); 2 June 1925, J. G. B. Beumeé 6767 (BO!); Cigelung-Dungusiwul, Jasinga, 23 Dec. 1926, J. G. B. Beumeé A.363 (BO!); Dungusiwul Jasinga, 15 May 1971, J. Dransfield JD 1507 (BO!); 15 May 1971, J. Dransfield JD 1508 (BO!, K!, L!); 15 Jan. 1998, A. Keim AK 30 (BO!); 16 Jan. 1998, A. Keim AK 31 (BO!); 20 June 1999, A. Keim AK 49 (BO!, K!). West Kalimantan: Anambas Islands, 1928, Henderson s.n. (K!); Karimata Islands Group, Pulau Penebangan, 20 March 1931, Mondi 120 (BO!); Gunung Palung, Cabang Panti Research Station, 26 July 1990, Palmiotto 1016 (BO!, K!). CULTIVATED. Java: Kebun Raya Bogor, 17 June 1999, A. Keim et al. AK 48 (BO!, K!). Singapore: Singapore Botanic Gardens, 8 Jan. 1933, Kiah 26132 (K!).

habitat. Lowland humid evergreen tropical rainforest from 0 to about 600 m above sea level. In some parts of Sumatra and Borneo sometimes found in heath forest (‘kerangas’).

conservation status. Least concern (LC). On a global scale this species is widespread and often abundant in, for example, Peninsular Malaysia and South Thailand. However, at the local level it may be seriously threatened; for example, in Java it is on the verge of extinction.

etymology. Growing in the forest.

vernacular names. ibul (Malay-Peninsula dialect), iwul (Sundanese), kayu baluhur (Malay-Asahan and Padang Sidempuan dialects), pon (Thai-Kao Panom dialect), kapun (Thai-Kao Salaw dialect), lee-boy (Thai-Thalae Ban dialect).

uses. Trunk is used for building houses. Leaves are used for house thatching. Fruits are said to be poisonous.

notes. Orania sylvicola is the second-most widespread species after O. palindan. This species is also known to inhabit a wide range of habitats, from humid lowland rainforests with rich soil to heath forests with very poor sandy soil. O. sylvicola is commonly found from coastal areas up to about 600 m above sea level. In Southern Thailand — around Tamben Kao Panom, Kerr 18802 — it has been reported from 1100 m. In Malay Peninsula O. sylvicola is found throughout the peninsula and on the adjacent island, Tioman. In Singapore O. sylvicola used to be found on mainland Singapore and a small offshore island (Ubin). However, this is based on reports and specimens collected by Ridley in 1891 (see Ridley 1900) at a time when a large part of Singapore was still covered with lowland tropical rainforest. The presence of O. sylvicola in Singapore and the surrounding islands has never been reported again. In Sumatra O. sylvicola is found throughout the island but mostly in the North and Central part of the island (i.e. the Provinces of North Sumatra and Jambi). Dransfield (1974) reported O. sylvicola in an area between Jambi and Bangko growing in lowland grassland (‘alang-alang’). In South Sumatra it has been reported growing in great numbers in heath forest (‘kerangas’) developed on very poor sandy soil just about 200 km from the coast (see Dransfield 1972). Apart from these two reports O. sylvicola normally grows in lowland forest on rich soil. So far no species of Orania has been reported from the northernmost part of the island (Aceh). As in Sumatra, in Borneo O. sylvicola is also found scattered throughout the island — mostly in the western part, but it is absent from Sabah. In Borneo O. sylvicola is also found in both lowland forest on well-drained soil (sometimes also in sandstone based soil as in West Kalimantan) and occasionally in heath forests (‘kerangas’) with less rich soil as in some parts of Sarawak (see Dransfield 1984). In Java O. sylvicola is restricted to the western part only. Nowadays this species can be seen only in the Dungus Iwul Nature Reserve, Jasinga, West Java. Another locality in West Java where O. sylvicola was once found is Pancoran Mas Nature Reserve-Depok (Backer & Bakhuizen van den Brink 1968). It was still there until at least the early 1970s (Dransfield 1998 pers. comm.). Unfortunately, when AK visited the place, O. sylvicola was no longer present. O. sylvicola has never been reported further east from West Java. Although there is another species that possesses the unequal staminodes, O. longistaminodia, several characters distinguish O. sylvicola from O. longistaminodia (Table 22).

Table 22 Morphological differences between Orania longistaminodia and O. sylvicola.

Distinct species not linked to other subgroups

There are four species, which cannot be placed in any of the subgroups previously described, namely Orania deflexa, O. gagavu, O. longistaminodia and O. zonae.

25. Orania deflexa A. P. Keim & J. Dransf. sp. nov.

Palma mediocris vel grandis, petiolo manifeste decurvo manifeste, foliolis in unum planum in medio rhachidis dispositis unum, basin irregulariter dispositis; inflorescentia 3-plo ramosa.Typus: Papua New Guinea, Morobe, near Bulolo, Hidden Valley, near Mount Kaindi, close to Wau, 07°19'1.7''S 146°37'26.5''E, 8 Sept. 2006, W. J. Baker 1319 (holotypus K!; isotypi BO!, LAE).

http://www.ipni.org/urn:lsid:ipni.org:names:77118916-1

Medium to large palm. Trunk to 10 m high, 20 cm diam. near the crown, internodes 2 – 3 cm. Leaves 11 in crown, spirally arranged, 330 cm long; leaf-sheath 80 cm long, 6 – 7 cm wide, densely covered with red-brown tomentum, margin disintegrating into fibres; petiole 90 cm long, 3 cm diam., strongly decurving (arching downwards) giving the crown drooping appearance, adaxial surface densely covered with wax and creamy brown tomentum, abaxial surface sparsely covered with wax; leaflets elongate-lanceolate, total 102, irregularly arranged in lower $\frac{2}{3}$ of rachis, in terminal $\frac{1}{3}$ rachis more or less regular, held in more than one plane in middle half of leaf giving at least $\frac{1}{3}$ a plumose appearance, the proximal 3 – 4 leaflets crowded 0.5 – 1 cm distant in a group, otherwise 7 cm distant in middle part of rachis, distance between 2 leaflets in the proximal part 1 cm, otherwise 5 – 7 cm distant in middle part of rachis, leaflets c. 90 – 95 cm long, 5 – 5.5 cm wide, adaxial surface glabrous, red-brown tomentum on midrib, midrib thick, other ribs slender, glabrous, abaxial surface densely covered with white indumentum and wax, midrib massive, other ribs glabrous and slender. Inflorescence spreading, branching to 3 orders, 180 cm long, densely covered with red-brown tomentum; prophyll persistent, disintegrating into fibres from top to base, 40 × 17 cm; peduncle 65 cm, covered with dense creamy brown tomentum; peduncular bract 1, woody, splitting in the middle, persistent, adaxial surface glabrous, abaxial surface densely covered with red-brown tomentum; rachis 115 cm long, covered with dense creamy white indumentum and red-brown tomentum; first order branches 75 cm long, densely covered with red-brown tomentum; second order branches 3 – 5 cm long, densely covered with red-brown tomentum; rachillae slender, straight, not conspicuously zigzag, fairly glabrous, 29 – 32 cm long, bearing 100 – 110 flower clusters, bearing arranged triads in the proximal $\frac{2}{3}$ and paired staminate flowers in the distal $\frac{1}{3}$ of rachilla, the basal 0.5 cm devoid of flowers, triads 1 cm distant, rachilla surface fairly glabrous. Staminate flowers with calyx of 3 united minute, scale-like sepals; corolla with 3 free petals, 7 – 8 × 2 – 3 mm; stamens 6, filaments free, dark brown, c. 1 mm long, anthers elongate-lanceolate, pale creamy-yellow, always free, c. 3 mm long each; pistillode absent. Pistillate flowers with calyx of 3 minute united sepals; corolla with 3 free petals, 6 × 4 mm, creamy yellow to yellow or orange when mature; staminodes 6, ½ as long as the gynoecium; gynoecium dark brown, 4 × 5 mm; stigma of 3 elongate lobes. Fruits unknown. Embryo unknown. Eophyll unknown. Fig. 9.

distribution. Endemic to Papua New Guinea. Map 9.

specimen examined. papua new guinea. Morobe: Near Bulolo, Hidden Valley, near Mount Kaindi, close to Wau, 07°19'1.7''S 146°37'26.5''E, 8 Sept. 2006, W. J. Baker 1319 (holotype K!; isotypes BO!, LAE).

habitat. Disturbed montane forest at altitude of 1400 m.

conservation status. Critically endangered (CR B1 B2). When the type collection was made in 2006, the population was estimated to be fewer than 100 individuals, growing in an area much disturbed by an active gold mine (W. J. Baker, pers. comm.).

etymology. Decurving (arching downwards), referring to the leaf orientation.

vernacular names. Unknown.

uses. Unknown.

notes. The decurving leaf orientation is so far unique to Orania deflexa, thus a very good character for field identification. O. deflexa is also known as one of the three Malesian species that possess the three branching orders inflorescences, the other two are O. gagavu and O. glauca. For differences see Table 20.

26. Orania gagavu Essig (1980: 227 – 228). Type: Papua New Guinea, Milne Bay, on slope of Mt Daraia, several km N of Kaporika Village, Alotau Sub-province, 12 May 1978, Essig & Young LAE 74096 (holotype LAE; isotypes BH!, L!, USF).

Large palm. Trunk to 7 m tall, c. 20 – 25 cm diam. (dbh). Leaves spirally arranged; leaf-sheath & petiole 1 m long, adaxial surface glabrous, abaxial surface densely covered with red-brown tomentum; petiole with red-brown tomentum; rachis c. 3 m long, c. 1.8 – 2.5 cm diam. in middle part, with dense red-brown tomentum; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, the proximal 3 leaflets crowded 2 – 3 cm distant in a group, otherwise c. 6.5 cm distant in middle part of rachis, c. 110 to 120 cm long, 4.5 – 5.5 cm wide; adaxial surface dark green, glabrous, red-brown tomentum on the midrib, midrib thick, other ribs slender, glabrous; abaxial surface densely covered with white indumentum, slightly red-brown tomentum on the margin, midrib thinner than adaxial, other ribs slender. Inflorescence spreading, branching to 3 orders, glabrous, c. 2.5 m long; peduncle c. 1 m long; rachis c. 1.5 m long; 36 first order branches, c. 94 – 95 cm long, secondary branches c. 65 – 70 cm long, rachillae 55 – 65 cm long, slender, distally zigzag, triads in the proximal $\frac{1}{2}$ – $\frac{3}{5}$, the basal devoid c. 3 – 4 cm of flowers, triads c. 2.5 – 3.5 cm distant, rachilla glabrous, number of flower clusters 80 – 90 in rachilla, triads in the proximal 20 – 25. Flowers unknown. Fruits globose, young fruit with c. 9 mm diam. Embryo placed below middle line of seed. Eophyll bifid.

distribution. Endemic to Papua New Guinea. Map 9.

specimens examined. papua new guinea. Milne Bay: On slope of Mt Daraia, several km N of Kaporika Village, Alotau Subprovince, 10°20'S 150°10'E, 12 May 1978, Essig & Young LAE 74096 (LAE holotype, BH!, L!, USF).

habitat. Rainforest, on mountain slopes, 400 m above sea level.

conservation status. Data deficient (DD). This palm is known only from its type and we have no recent information on the status of forests at the type locality.

etymology. Gagavu, local name.

vernacular names. gagavu (Kaporika).

uses. Unknown.

notes. Apart from the number of orders of inflorescence branching and flower clusters in a rachilla (Table 23), Orania gagavu closely resembles O. lauterbachiana. Although it is not common in the genus, the variation in the number of inflorescence branching orders is not unusual in Orania (e.g. O. paraguanensis and O. trispatha). Unfortunately, we only have one collection of O. gagavu. Nevertheless, until we have more specimens, we regard the two species (O. gagavu and O. lauterbachiana) as distinct based on two characters (Table 23).

Table 23 Morphological differences between Orania gagavu and O. lauterbachiana.

27. Orania longistaminodia A. P. Keim & J. Dransf. sp. nov.

Palma grandis, a ceteris speciebus floribus staminibus 5 ad 7, staminodiis florum pistillatorum inaequalibus, grandibus 2 staminodiis similibus (c. 5 mm longis unusquisque) vel 3 staminodiis inaequalibus – uno 2 ad 2.5 mm longo, duo c. 5 mm longo, differt. Typus: Papua New Guinea, East Sepik, near Antenna, 16 Nov. 1996, Barfod et al. 374 (holotypus K!; isotypi AAU, LAE).

http://www.ipni.org/urn:lsid:ipni.org:names:77118917-1

Large palm. Trunk c. 6 m tall, c. 22 cm diam. (dbh), internodes 5 – 10 cm, nodes c. 3 – 5 cm wide. Leaves 12 in the crown, spirally arranged; leaf-sheath and petiole 2.6 – 2.8 m long, adaxial surface glabrous, abaxial surface with thin red-brown tomentum, margins disintegrating into fibres; petiole c. 4 – 5 cm diam., margins disintegrating into fibres, with thin red-brown tomentum; rachis c. 3.63 m long, covered with thin red-indumentum; leaflets elongate-lanceolate, regularly arranged leaflets held in one plane, the proximal part of rachis with 1 single leaflet, c. 8.5 cm distant in the middle part of rachis, c. 130 × 10 cm, adaxial surface glabrous, with thin red-brown tomentum on the midrib, midrib thick, other ribs slender, glabrous, abaxial surface densely covered with white indumentum, red-brown tomentum on the margin and basal part of the leaflet, midrib robust, other ribs slender. Inflorescence spreading, branching to 2 orders, c. 53 – 94 cm long; prophyll persistent, heavily disintegrating into fibres, c. 33.5 – 34 cm long, 9 cm wide, distally splitting in the middle; peduncle c. 33 – 58 cm long, with red-brown tomentum; peduncular bract one, woody, adaxial surface glabrous, abaxial surface with red-brown tomentum, c. 80 – 90 cm long, persistent, splitting in the middle, disintegrating into fibres; rachis c. 20 – 36 cm long; first order branch c. 25 cm long, rachillae bract c. 9 × 20 mm; rachillae thick, c. 18 – 20 cm long, bearing 36 – 42 flower clusters, triads arranged in the proximal $\frac{1}{3}$ & staminate flowers in the distal $\frac{2}{3}$ of rachilla, the basal c. 1.5 – 2.5 cm devoid of flowers, triads c. 1.5 cm distant, maximum number of pistillate flowers in rachilla 8. Staminate flowers with calyx of 3 united minute sepals; corolla with 3 free petals, c. 5 × 2 mm; stamens 5 – 7, filaments free, dark-brown, 0.7 – 1 mm long, anthers elongate-lanceolate, pale creamy yellow, free, c. 2 – 3 mm long, pistillode uncommon, c. 2 – 2.5 mm long, 3 – 3.5 mm wide. Pistillate flowers with calyx of 3 united sepals, c. 2 mm long; corolla with 3 free petals, c. 8 × 6 mm; staminodes 6, unequal, 3 small, c. 1 mm long, 1 large, c. 2 – 2.5 mm long, 2 conspicuously large, c. 5 × 2 mm, in some flowers with 3 conspicuously large, c. 5 × 2 mm and 3 small staminodes, c. 1 mm long; gynoecium dark-brown, c. 4 mm long, 4 – 5 mm wide; stigma with 3 elongate lobes, c. 1 – 1.5 mm long. Fruits globose or bilobed, c. 7 cm diam., otherwise c. 4.8 – 5 cm diam. (Barfod 473), bright orange when mature, stigmatic remains sub-basal (Barfod et al. 374). Embryo placed below middle line of seed. Eophyll bifid. Fig. 10.

distribution. So far this species has been collected only from the eastern part of New Guinea (i.e. Papua New Guinea) with a quite disjunct distribution. One specimen was collected from the northern part of the country, whereas the other one was from the southern part. Map 9.

specimens examined. papua new guinea. East Sepik: Near Antenna, 143°37'S 3°37'E, 16 Nov. 1996, Barfod et al. 374 (K! holotype, AAU, LAE). Gulf: Logging camp on the Vailala R., 145°29'S 07°45'E, 8 – 9 March 2000, Barfod et al. 473 (AAU, BRI, CANB, K!, LAE).

habitat. Disturbed lowland tropical rainforest in coastal area at low elevation.

conservation status. Data deficient (DD). The palm is known from only two specimens from widely separated localities. We have no information on the status of forests at these localities.

etymology. Long staminodes.

vernacular names. Unknown.

uses. Sap is said to be used by local tribes as a medicine for sore throats (Barfod et al. 374, field notes).

28. Orania zonae A. P. Keim & J. Dransf. sp. nov. Palma grandis, a ceteris speciebus inflorescentia expansa, staminibus 6, 9 vel 12, antheris liberis (in staminis 6), vel connatis in staminis 9 connatis, vel antheris omnibus connatis, filamentis conatis, staminodiis 6 differt. Typus: Indonesia, Papua Barat, Manokwari, Sungai Asai between Mt Manggombo and Marwadibau, 16 Aug. 1995, Zona SZ 674 (holotypus K!; isotypi BO, FTG, MAN).

http://www.ipni.org/urn:lsid:ipni.org:names:77118918-1

Large palm. Trunk 20 m tall, c. 27.2 cm diam. (dbh). Leaves 12 in the crown, spirally arranged, c. 4.3 m long; leaf-sheath c. 45 cm long, 7 – 8 cm wide, margins disintegrating into fibres, curly, c. 20 cm long, adaxial surface glabrous, abaxial surface with red-brown tomentum; petiole c. 2 m long, with red-brown tomentum, margins massively disintegrating into a dense fibrous mass; rachis c. 1.85 m long, c. 3 cm diam. in the middle, with red-brown tomentum; leaflets elongate-lanceolate, 61 pairs totally, regularly arranged leaflets held in one plane, leaflets c. 9 cm distant, c. 1.47 – 1.9 m long, 8.5 – 11 cm wide, adaxial surface glabrous, with thin red-brown tomentum on midrib, midrib robust, other ribs less robust; abaxial surface with dense white indumentum, dense red-brown tomentum on the margin, thin on the midrib, midrib robust, other ribs less thick than adaxial. Inflorescence spreading, robust, branching to 2 orders; prophyll persistent, c. 83.5 – 85 cm long, 12 cm wide, adaxial surface glabrous, abaxial surface with dense red-brown tomentum, massively disintegrating into fibres when old; peduncle with dense red-brown tomentum; peduncular bract one, woody, persistent, abaxial surface glabrous, adaxial surface densely covered with red-brown tomentum, c. 1.4 m long, splitting in the middle, disintegrating into fibres when old; rachis c. 2 – 3 cm diam., with dense red-brown tomentum; first order branches c. 109 – 110 cm long; rachillae thick, c. 45 – 50 cm long, bearing 90 – 100 flower clusters, bearing triads in the proximal $\frac{3}{5}$, the basal c. 2 cm devoid of flowers, triads c. 1 cm distant. Staminate flowers white; with calyx of 3 united minute sepals; corolla with 3 free petals, 3 × 2 mm; stamens 6, 9 or 12, filaments united, dark-brown, c. 0.5 mm long, anthers elongate-lanceolate, pale creamy yellow, c. 1 mm long, in stamens 6 or 9 anthers free, in stamens 12 anthers united in 6 pairs; pistillodes absent. Pistillate flowers white; with calyx of 3 united minute sepals; corolla with 3 free petals, c. 4 × 3.5 mm; staminodes 6, c. 1.5 mm long; gynoecium dark-brown, c. 3 mm long, stigma of 3 elongate lobes, c. 1 mm long, brighter coloured. Fruits globose or bilobed, c. 4.0 cm diam., dull green when young, red when mature (Zona 1995 pers. comm.). Embryo placed below middle line of seed. Eophyll bifid. Fig. 11.

distribution. Endemic to an area near the Asai River in the district of Manokwari — formerly Vogelkop in the Indonesian province of Papua Barat. Map 9.

specimen examined. indonesia. Papua Barat. Manokwari: Sungai Asai between Mt Manggombo and Marwadibau (133°56.5'E by 0°45'S), 16 Aug. 1995, Zona et al. SZ 674 (K! holotype, BO, FTG, MAN).

habitat. Undisturbed lowland forest on limestone hill with elevation about 165 m above sea level.

conservation status. Data deficient (DD). This palm is known only from its type. APK searched for it but without success.

etymology. After Scott Zona, American palm taxonomist and collector of the type specimen.

vernacular names. motuaga (Meyah language).

uses. Outer wood used for arrowheads (Zona 1995 pers. comm.).

notes. An attempt had been made to recollect this species from the type locality in 1999, without success.

Doubtful & excluded species

Orania appendiculata (F. M. Bailey) Domin (1915: 498). Type: Areca appendiculata F. M. Bailey.

Areca appendiculata F. M. Bailey (1891: 18). Type: Australia, Queensland, Bellenden-Ker, F. M. Bailey s.n. 1889 (holotype BRI).

Orania beccarii F. M. Bailey (1909: 35). Type: Areca appendiculata F. M. Bailey.

notes. For a full discussion of this taxon and its transfer to a separate genus, Oraniopsis, as Oraniopsis appendiculata (F. M. Bailey), J. Dransf. et al. belonging to a different palm subfamily (Ceroxyloideae) see Dransfield et al. (1985).

Orania nicobarica Kurz (1875: 331). Type: India, Nicobar Island, Kurz s.n. (holotype CAL).

notes. This species was included by Beccari (1885) as a synonym for a species belonging to a different genus, Bentinckia nicobarica Becc.

Orania nivea Linden ex W. Watson (1887: 157). nom. nud.

notes. This is a name based on a juvenile plant cultivated in Kew with no adequate description and herbarium material. The name is thus regarded as being of uncertain application.

Orania porphyrocarpa Blume in Martius (1849: 190, t.157).

notes. The first record of this taxon found in Java was by Blume (1836) under the name Orania regalis Blume (Rumphia 2, plate 95) and for a few years had been confused with O. regalis Zipp. Martius then reassessed the species and put it under the name O. porphyrocarpa (in Historia Naturalis Palmarum 3: 187. 1838 – 1839 (ed. 1) and page 190 in 1849 (ed. 2)). Moore (1960) reassessed the species again and transferred it to Arenga as A. porphyrocarpa (Blume) H. E. Moore.