The genus Pandanus Parkinson (Pandanaceae) on Halmahera Island (Moluccas, Indonesia) with descriptions of three new species and a key to the species on the island

© CONSERVATOIRE ET JARDIN BOTANIQUES DE GENÈVE 2015 ISSN : 0373-2967 – Online ISSN : 2235-3658 – Candollea 70(2) : 179-195 (2015) The genus Pandanus Parkinson (Pandanaceae) on Halmahera Island (Moluccas, Indonesia) with descriptions of three new species and a key to the species on the island Martin W. Callmander, Ary P. Keim, Sven Buerki & Peter B. Phillipson Addresses of the authors : MWC: Missouri Botanical Garden, P.O. Box 299, St. Louis, MO, 63166-0299, U.S.A. Currently at: Conservatoire et Jardin botaniques de la Ville de Genève, C.P. 60, 1292 Chambésy, Switzerland. E-mail: martin.cal[email protected] APK: Botany Division/Herbarium Bogoriense (BO), Centre Research for Biology, P2B LIPI, Jl. Raya Jakarta-Bogor Km 46 Cibinong, Bogor 16911, West Java, Indonesia. SB: Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom. PBP: Missouri Botanical Garden, P.O. Box 299, St. Louis, MO, 63166-0299, U.S.A. and Institut de Systématique, Évolution, et Biodiversité (UMR 7205 – CNRS MNHN EPHE UPMC), Muséum national d’Histoire naturelle, C.P. 39, rue Cuvier 57, 75231 Paris CEDEX 05, France. Submitted on March 16, 2015. Accepted on April 6, 2015. Edited by P. Bungener Abstract CALLMANDER, M. W., A. P. KEIM, S. BUERKI & P. B. PHILLIPSON (2015). The genus Pandanus Parkinson (Pandanaceae) on Halmahera Island (Moluccas, Indonesia) with descriptions of three new species and a key to the species on the island. Candollea 70: 179-195. In English, English abstract. DOI: http://dx.doi.org/10.15553/c2015v702a2 Halmahera is the largest (c. 18,000 km2) island of the Moluccan archipelago, but naturalists have only sporadically visited Halmahera and it has remained very poorly explored botanically. However, an intensive botanical inventory project was undertaken between 2012 and 2014 in part of the island to inform flora biodiversity management for certain proposed mining activities. This effort has contributed over 3600 plant collections and nearly doubled the number of Pandanus Parkinson (Pandanaceae) specimens (bringing the total to 55) available for Halmahera. After careful examination of all available material and comparison with other material from the region, we are able to present the first overview of the genus for the island. We have identified ten species from the island of which three are new to science and not known elsewhere, while the other seven are all representatives of species already described from other localities. The new species are formally described here as Pandanus beguinii Callm. & A.P.Keim, Pandanus benstoneoides Callm., Buerki & Phillipson and Pandanus halmaherensis Callm. & A. P. Keim. The new species are provided with notes on their respective morphology and known distributional and ecological ranges, line drawings. Those three new species are assigned a preliminary status of Endangered following IUCN Red List Categories and Criteria. All ten species are illustrated with colour photographs and a key to the species is provided. Keywords PANDANACEAE – Pandanus – Moluccas – Halmahera – Taxonomy – New Species – IUCN Red List 180 –The genus Pandanus (Pandanaceae) on Halmahera Island (Moluccas, Indonesia) Candollea 70, 2015 Introduction Halmahera is the largest (c. 18,000 km2) island within an archipelago of approximately 2896 islands in Indonesia collectively known as the Moluccas. Administratively the Moluccas are now divided into two provinces: North Maluku, which includes Halmahera, and Maluku, which includes the islands of Ambon and Seram to the south. The Moluccas, together with Sulawesi to the east and a long chain of islands to the south, form the biogeographic region known as Wallacea (Fig.1), which is a biodiversity hotspot lying between Borneo and New Guinea (Myers et al., 2000). After Sulawesi, Halmahera is the second largest island of Wallacea. Halmahera has a complex geological history situated at the interference of tectonic plates associated with past and active volcanic arcs (see Hall et al., 1988). The geology of the island consisting now of a mixture of sedimentary rocks resulting from raised coral reefs, intrusive igneous rocks and fragments of continental crust (see Hall, 2001). The coastline of Halmahera is just 13.5kilometres away from the small, but economically dominant island of Ternate, one of the so-called “Spice Islands”, which is particularly known for the cultivation of cloves (Ricklefs, 2008). Historically, botanical exploration ofthe region focused on Ternate, while the flora of Halmahera, despite it being the largest island of the Moluccas, was largely ignored and has remained very poorly known. Among the more extensive collections from the island were those made by Victor E. Beguin, a Dutch planter based in Java, who was hired by the Museum and Inquiry Office for Economic Botany in Buitenzorg (now Bogor) to make extensive travels to the Moluccas (van Steenis, 1950). He visited different parts of Halmahera (see Backer, 1936) and Ternate between 1920 and 1923 and left a legacy of more than 1750herbarium collections from these travels, including 640 from Halmahera (van Steenis, 1950). During the second half of the XXth Century only a handful of botanists have made more than a hundred collections each on Halmahera, but among these the contribution of Eduard (“Ed”) de Vogel (682 collections) is certainly the most important. The better-known fauna of Halmahera is however characterized by exceptional species-level endemism but is only moderately diverse (Setiadi et al., 2010). Rumphius was the first to publish some information about the flora of Halmahera (then known by its indigenous name “Gelolo”) in his legendary six volumes series of “Herbarium Amboinense” (Rumphius, 1741-1755). However, Rumphius probably never visited Halmahera himself despite his numerous visits to other islands adjacent to Ambon where he was based. Rumphius first mentioned Pandanus ceramicus Rumph. [nom. inval.] (= P. conoideus Lam.) as being present on Halmahera, presumably on the basis of reported observations. The earliest Pandanaceae specimens known from Halmahera were collected by Johannes E. Teijsmann and Willem H. de Vriese in their “Herb. Itineris in Insulas Moluccanas” series in 1859 (van Steenis, 1950). They twice collected P. polycephalus Lam. (Vriese & Teijsmann 18; Teijsmann 87; see Table 1), when they briefly visited Halmahera in the vicinity of Sidangolie (now Sidangoligam) on the coast facing Ternate (Teijsmann, 1861). Surprisingly, the Italian naturalist Odoardo Beccari, who travelled extensively in the region between 1870 and 1875, never visited Halmahera (Beccari, 1924). No other records of Pandanaceae are known from Halmahera prior to the collections of Victor Beguin, who clearly had an interest in the family and made ca. 50 collections of Pandanaceae from Halmahera. Afew additional Pandanus Parkinson collections were made during the second half of the XXth and early XXIst Century, notably by several ethnologists who visited Halmahera for their studies (Yoshida, 1980; Taylor, 1990; Purwanto, 2009). In 2012, the Missouri Botanical Garden, in collaboration with Herbarium Bogoriense and the National Herbarium of the Netherlands, was engaged by Weda Bay Nickel (WBN) to undertake a botanical inventory on the company’s Contract of Work (CoW), an extensive area spanning the Central and Eastern Halmahera Regencies (Fig. 1) to inform the process of flora biodiversity management in relation to their proposed mining project. This work built on earlier baseline studies undertaken for an impact assessment, which resulted in the identification of several Pandanaceae species, but unfortunately no voucher specimens were preserved. Between September 2012 and December 2014, a locally-based inventory team was trained and more than 3600 botanical collections were made on the CoW, and critical identification of the material is being undertaken by specialists. The collections include 65Pandanaceae in three genera: Benstonea Callm. & Buerki [1 collection], Freycinetia Gaudich. [39 collections] and Pandanus [25 collections]. Benstonea is morphologically distinct by its one-seeded drupes with its stigma always positioned on the abaxial side of the its acutely-pointed, linear style. This genus is represented by only a single species: B. verruculosa (B. C.Stone) Callm., Buerki & Phillipson, which is endemic to Halmahera and which was the subject of a previous publication (Callmander et al., 2014). Freycinetia, on the other hand, is characterized by its lianescent habit, multi-ovulate carpels and berry fruits and will be the subject of a forthcoming publication on the diversity of the genus on Halmahera. Callmander et al. (2012) provide a key to the genera of Pandanaceae and Buerki et al. (2012) and Gallaher et al. (2015) report on molecular phylogenetic studies for the family. The 25 newly collected Pandanus specimens together with Beguin’s legacy of 25 Pandanus specimens and the five other available collections allow us to present here a first overview of the genus for Halmahera. In this article we document 10species of Pandanus that we have identified from Halmahera. These include seven species that are already known to Candollea 70, 2015 The genus Pandanus (Pandanaceae) on Halmahera Island (Moluccas, Indonesia)– 181 science and also occur in other parts of South-East Asia, and for which we provide descriptive notes. In addition, our studies have revealed the presence of three species that are new to science and which are, based of current knowledge, endemic to Halmahera. In this article we formally describe the three new species: P. beguinii Callm. & A. P. Keim, P. benstoneoides Callm., Buerki & Phillipson and P. halmaherensis Callm. & A.P. Keim and each of them are provided with notes on their morphology, relationships, conservation assessments and line drawings. We also provide an identification key to the ten species currently known from Halmahera, together with colour photographs for each species (Fig. 2, 3). The genus Pandanus in Halmahera The coastal P. dubius Spreng., P. polycephalus and P. tectorius Parkinson are widespread in littoral and lowland forest in swampy or sandy ground. Pandanus dubius is distributed from the Andaman Sea (Andaman Nicobar islands) to the western Pacific region (Kiribati, Federated states of Micronesia) and throughout Malesia. This ocean-dispersed species (Gallaher et al., 2015) is frequently cultivated in the tropics and the leaves are used for basketry and roof thatching. The albumen of the ripe fruit is edible and used in Micronesia as a source of starch (Lim, 2012). It is easily recognizable by its habit (large tree with robust proproots at the base), its wide, coriaceous leaves (11-16 × 200cm) with conspicuous longaristate apices and its large pendulous subspherical syncarp (20-30cm in diam.), composed of pale bluish-green drupes (Fig. 3D). Pandanus tectorius, another well-known, ocean dispersed and frequently cultivated species has a broad distribution from Malesia to the Southern Pacific islands (Fig.2E). This species, which is known to have various medicinal, nutritive and construction properties (Lim, 2012), is taxonomically problematic and is currently regarded as a species complex in need of further study (Gallaher et al., 2015). Pandanus polycephalus is also a strand plant on seashores, it reaches its western-most distribution in the Moluccas and is found through New Guinea and as far east as the Solomon Islands. The species forms a slender treelet to 10 m tall and can be easily recognized by its small multispicate infructescence with the drupes becoming red on ripening (Fig. 2H). Afourth species is confined to coastal areas, P. kaernbachii Warb. It is a robust solitary tree reaching up to 10 m high and can be easily identified in the field by its spicate infructescence consisting of 2 to 4 syncarps with incompletely fused phalanges (Fig.3I). Unlike P.tectorius, P. kaernbachii is always solitary or in scattered populations (Keim et al., 2009; Callmander, pers. obs.). The presence of this species in the Moluccas was firstly reported by Keim et al. (2008) from Seram Island, 200 km south of Halmahera. Pandanus krauelianus K. Schum. seems to be a common inland species at low to mid elevation on Halmahera (Table 1). It is part of the morphologically homogenous Pandanus sect. Maysops H. St. John, in subgen. Lophostigma (Brongn.) H. St. John, characterised by a solitary, ovoid-ellipsoid syncarp hidden by persistent bracts, one-seeded drupes and a pileus generally topped apically with a small hardened turret bearing a lateral stigma (Fig. 2F, 2G) (see also Stone, 1974a). Stone (1992) published a review of the section for New Guinea, and accepted ca. 16 species and emphasised the need of further collections in this group in which species delimitation is difficult. In the latter revision, Pandanus amboinensis Warb., the first valid publication of P.sylvestris Rumph. from Ambon, is considered to be different from P.krauelianus from New Guinea (West Papua, Indonesia). Jebb (1992) and Keim (2009) did consider these two species as synonyms and this broader species concept is followed here. Pandanus krauelianus, thus delimited, is distributed from the Moluccas to the Bismark archipelago through New Guinea. Another species with a similar distribution is 16° 127°E 128°E 129°E O° 1°N 2°N 60 km Fig. 1. – General Map of Halmahera Island and its localisation : west of New Guinea and east of Sulawesi (framed). 182 –The genus Pandanus (Pandanaceae) on Halmahera Island (Moluccas, Indonesia) Candollea 70, 2015 found in inland swamps at low elevation: P. papuanus Solms. It is a large tree species (to 20 m tall) with a large, impressive cone of proproots at the base and massive pendent syncarps (Fig. 3G, 3H). It is known from similar habitats at higher altitudes (ca.900m) in New Guinea (Stone, 1982). Pandanus conoideus, first mentioned by Rumphius (1741-1755), has not yet been collected on Halmahera, but a photograph of it taken by one of us (PBP) during his first visit to the island in 2011 confirmed its presence on the island (Fig. 3J). Pandanus conoideus is widely cultivated in New Guinea for its edible fruits and may have originated from the Moluccas (Stone, 1982). The 55 Pandanus collections currently known from Halmahera are summarized in Table 1. Key to the Pandanus species on Halmahera 1. Carpels free in one-carpellate and one-seeded drupes (rarely bicarpellate in P. dubius Spreng.) ............ 2 1a. Carpels connate into multicarpellate, several-seeded drupes ..................................... 8 2. Stigma filiform or spinescent and linear ........... 3 2a Stigma oblique, flat or on an apical turret, but not filiform or linear .................................... 4 3. Infructescence lateral, born just below the base of leaves; drupes with a yellow pileus; stigma filiform, appressed towards the apex of the syncarp, 15-20mm long (Fig.2A-D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. benstoneiodes Callm., Buerki & Phillipson 3a Infructescence terminal, pendant; drupes with a red to purple shiny pileus; stigma spinescent, sharp, erect, 6-8mm long (Fig. 3E, 3F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. halmaherensis Callm. & A. P. Keim 4. Syncarps spicately arranged, small (4cm long), ovoid; stigma reniform and flat (Fig. 2H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. polycephalus Lam. 4a Syncarp solitary, much larger (10-40cm long), ellipsoid to oblong; stigma with various shapes but not reniform and flat ........................................ 5 5. Leaves wide (11-16cm) ; drupes large (> 10cm) at maturity (Fig. 3D) .....................P. dubius Spreng. 5a. Leaves narrow (< 8cm) ; drupes smaller (< 8cm) at maturity ........................................ 6 6. Small monocaulous shrub (3 m tall) with few slender proproots at the base; leaves (<110cm long) ; and syncarp small (< 15cm) ; peduncle slender and short, c. 11cm long (Fig.2A-C) ......... P. beguinii Callm. & A. P. Keim 6a. Tall ramified tree (8-15 m) with large cone of proproots at the base; leaves (>180cm long) ; and syncarp large (>25cm) ; peduncle stout and long, > 30cm long .... 7 7. Syncarp yellow at maturity; pileus topped apically with a small hardened turret bearing a lateral stigma (Fig. 2F, 2G) ..................... P. krauelianus K. Schum. 7a. Syncarp red at maturity; pileus with central vertical stigma (Fig. 3J) ........................P. conoideus Lam. 8. Syncarps spicately arranged (Fig. 3I) .............. ...........................P. kaernbachii Warb. 8a. Syncarp solitary .............................. 9 9. Large tree (to 20 m tall) with a large and impressive cone of proproots at the base; leaves large (200-600cm × 10-15cm) ; stigmas borne on the inner edge of the truncate carpel apices (Fig. 3G, 3H) ........P. papuanus Solms 9a. Medium-sized tree (< 15 m tall) with a narrow cone of proproots at the base; leaves smaller (mostly 60-150 × 3-6cm) ; stigma borne on the top of each of the rounded carpel apices (Fig. 2E) ..........P. tectorius Parkinson Systematics Pandanus beguinii Callm. & A. P. Keim, spec. nova (Fig.3A-C, 4A, 5). Typus: indoneSia. Prov. North Maluku [Halmahera Isl.] : Central, Weda Bay, near Weda Bay, 00°28’38”N 127°59’38”E, 26 m, 30.I.2013, Callmander, Haris, Lasut & Mahroji 1088 (holo- : G [G00341586] ! ; iso- : BO!, L!, MO-6465094!, MO-6465095!). Haec species a congeneris habitu gracili, syncarpo parvo terminali erecto drupas truncatas in turriculam obscuram conicam lateraliter concavam stigma minutum laterale bicornutum gerentem desinente atque inflorescentia staminata ex spica solitaria constante distinguitur. Shrub to 3 m tall, monocaulous; stem c. 3cm in diam., with few proproots at the base. Leaves spirally arranged, linear, mostly (50-)70-110cm long, (4-)5-8cm wide in the middle, but gradually narrowing to 2-3cm wide and more deeply folded in the lower third above the broader sheath, apex abruptly attenuate with a short acumen (3-10mm), chartaceous; longitudinal veins visible and prominent on both surfaces; marginal prickles borne 3-4cm above base to apex, antrorse, 0.4-1mm long in the lower third, 0.5-1mm apart, 0.7-1mm long in the mid-third, 4-7mm apart, to 0.5-1mm long in the distal third, 0.5-1.5mm apart, subappressed to appressed in the distal third; midrib unarmed, except the distal 7-13cm with prickles > 1.5mm long towards the apex, irregularly spaced but mostly 1-5mm apart; sometimes sparsely armed on apical ventral pleats; sheath (2.5-)4-4.5cm long, (2-)3-4cm wide at apex, (3-)4-6cm at base. Infructescence terminal, erect, green-orange when immature becoming red at maturity, the solitary syncarp ellipsoid, 10-15cm long, 5-6cm wide, peduncle short and slender, c. 11cm long, c. 1cm thick at apex, bearing multiple foliaceous bracts, crowded at peduncle apex, partly covering the syncarp, the proximal ones c. 45cm long, the distal one 15cm long. Drupes 400-500, 13-15mm long, 5-7mm wide, 4-5mm thick, 1-celled, (4-)6-angled, ovoid, truncate, pileus Candollea 70, 2015 The genus Pandanus (Pandanaceae) on Halmahera Island (Moluccas, Indonesia)– 183 Fig. 2. – Field pictures of Halmahera Pandanus Parkinson species. A, D. Pandanus benstoneoides Callm., Buerki & Phillipson (pistillate plant) ; B, C. Pandanus benstoneoides Callm., Buerki & Phillipson (staminate plant) ; E. Pandanus tectorius Parkinson; F, G. Pandanus krauelianus K. Schum. ; H. Pandanus polycephalus Lam. [Photos: A: E. Bidault; D, F-G: M. Callmander; B-C, E: P. Phillipson; H: M. Merello] A C F D G E H B 184 –The genus Pandanus (Pandanaceae) on Halmahera Island (Moluccas, Indonesia) Candollea 70, 2015 Fig. 3. – Field pictures of Halmahera Pandanus Parkinson species. A, C. Pandanus beguinii Callm. & A. P. Keim (pistillate plant) ; B. Pandanus beguinii Callm. & A. P. Keim (staminate plant) ; D. Pandanus dubius Spreng. ; E, F. Pandanus halmaherensis Callm. & A. P. Keim; G, H. Pandanus papuanus Solms; I. Pandanus kaernbachii Warb. [photo taken in Seram] ; J. Pandanus conoideus Lam. [Photos: A: M. Merello; B-G: M. Callmander; H, J: P. Phillipson; I: A. Keim] A D G H I J E F B C Candollea 70, 2015 The genus Pandanus (Pandanaceae) on Halmahera Island (Moluccas, Indonesia)– 185 Table 1. – The 55 Pandanus Parkinson collections currently known from Halmahera listed by species. Species Name Senior collector N° Locality Alt. [m] Date Herbaria Pandanus beguinii Beguin 1819 Loleo Poeloe Rao 13 Oct. 1921 BO Beguin 1822 Loleo Poeloe Rao 30 14 Oct. 1921 BO Beguin 1825 Loleo Poeloe Rao 40 15 Oct. 1921 BO Beguin 2044 Soa Tobaroe 60 24 June 1922 BO Beguin 2067 Soa Tobaroe 60 28 June 1922 BO Beguin 2069 Soa Tobaroe 60 29 June 1922 BO, L Beguin 2071 Soa Tobaroe 60 29 June 1922 BO Beguin 2107 Sao Tobaroe 60 29 June 1922 BO, L Beguin 2167 Soa Tobaroe 60 6 Aug. 1922 BO Bidault 1089 Weda Bay, on the road to Bukit Limber 600 15 Apr. 2013 BO, G, L, MO Callmander 1086 Near Weda Bay 15 29 Jan. 2013 BO, G, L, MO Callmander 1088 Sagea Lagoon 26 30 Jan. 2013 BO, G, L, MO Callmander 1090 Pinto 340 01 Feb. 2013 BO, G, L, MO Merello 3539A Kao Rahai 366 09 Dec. 2012 BO, MO Pandanus benstoneoides Bidault 1184 Bukit Limber 942 30 Apr. 2013 BO, G, L, MO Callmander 1077 Km 3 from Lelilef on the road to Doromesmesan 70 26 Jan. 2013 BO, G, L, MO, PH Callmander 1096 Pinto 340 1 Feb. 2013 BM, BO, G, L, MO Gushilman 646 Bukit Limber 1031 22 May 2013 BO, G, K, MO Gushilman 739 Tarsan Camp 495 2 June 2013 BO, G, L, MO Phillipson 6488 On road to Bukit Limber 712 12 Sept. 2013 BO, G, L, MO Phillipson 6489 On road to Bukit Limber 711 12 Sept. 2013 BO, G, K, L, MO Pandanus dubius Beguin 1770 Galela 1 24 Sep. 1921 BO, L Beguin 1841 Galela 3 24 Oct. 1921 BO, L Beguin 1842 Galela 3 24 Oct. 1921 L Callmander 1081 Along road to Gemaf village 527 Jan. 2013 BO, G, L, MO Pandanus halmaherensis Bangun 843 Sake West 218 19 June 2013 BO, G, L, MO Callmander 1078 along the Road to Bukit Limber 600 27 Jan. 2013 BO, G, L, MO Callmander 1114 SE of Tofu Blewen camp 480 03 Feb. 2013 BO, G, K, L, MO Fabanyo 58 Kao Rahai 693 26 Feb. 2013 BO, G, L, MO Gushilman 560 Weda Bay 163 11 Apr. 2013 BO, MO Pandanus kaernbachii Beguin 1812 Loleo Poeloe Rao 1922 [?] BO, L Beguin 1815 Loleo Poeloe Rao 60 12 Oct. 1921 L Beguin 1962 Tobelo 4 16 May 1922 BO, L Beguin 2214 Foeteo[ ?] ma Loleo 27 Sept. 1922 BO, L Beguin 2281 Tobelo 2 13 Dec. 1922 BO Beguin 2314 Berg Doekono, W Tobelo 800 28 Dec. 1922 BO, L 186 –The genus Pandanus (Pandanaceae) on Halmahera Island (Moluccas, Indonesia) Candollea 70, 2015 short, truncate, 2-3mm long, with a flat to slightly concave dark coloured turret, bearing a lateral minute bicornate stigma, c. 0.5mm wide. Endocarp bony, median, c. 5mm long, circular, lateral walls > 0.5mm thick; proximal mesocarp c. 2-3mm long, fibrous, distal mesocarp c. 4-5mm long, fibrous, chambered, seed locule c. 4mm, spherical. Staminate inflorescence c.30cm long with a single terminal spike on a fleshy peduncle, c. 20cm, covered with c. 9 caniculate foliaceous yellow bracts, distal ones c. 25 × 3.5cm; proximal ones 8 × 1cm; spike brown, short cylindric, mostly 11cm long, 2cm in diam., composed of crowded and congested flowers, formed by 4-12 briefly basally connate stamens; stamens c. 3mm long; with a dilated basal column, c. 2 × 0.5mm, the single anther born on a minute filament, c. 0.3mm long; anther c. 1 × 0.3mm, apiculate; apiculus c. 0.1mm long. Etymology. – This new species honours Victor E. Beguin (1886-1943) who collected this species nine times during his visits to Halmahera in 1921 and 1922 among the 50 collections of Pandanaceae gathered from the island. Distribution and ecology. – Pandanus beguinii is mostly found at lower elevations (30-600 m) in primary and disturbed evergreen tropical forest. It grows on slopes and in open areas, where individuals are scattered and discrete. The species is known from the northern, eastern and central regions of Halmahera (Fig. 4A). Conservation status. – With an EOO of 1625 km2, an AOO of 54 km2, and 5 subpopulations, none of which are situated within the North Maluku protected area network, P. beguinii is assigned a preliminary status of “Endangered” [EN B1ab(iii)+2ab(iii)] based on the IUCN Red List Categories and Criteria (IUCN, 2012) (calculation following Callmander et al., 2007). Four of the known subpopulations of P. beguinii have been recorded on or near WBN’s CoW, and we assume that some or all of them will be impacted by the proposed mining operations or associated environmental degradation. The current status of the subpopulation sampled by Beguin in northern Halmahera nearly a hundred years ago is unknown, but it occurs in an area that appears to be already somewhat degraded as judged by available satellite imagery. The conservation status is based on the currently available information on the geographical distribution of the species, and is justified because the extent of suitable habitat is expected to decrease in the coming years. Nevertheless, Halmahera is botanically poorly known and new subpopulations may occur in currently unexplored areas. If new Species Name Senior collector N° Locality Alt. [m] Date Herbaria Beguin 2315 Berg Soekono, W Tobelo 800 28 Dec. 1922 BO Callmander 1076 Doromesmesan 70 26 Jan. 2013 BO, G, L, MO Callmander 1132 NE of Tofu Blewen camp 610 05 Feb. 2013 BO, G, L, MO Idjan 303 Gunung Sombilan s.d. BO, L Pandanus papuanus Beguin 1372 Tobelo 10 26 Jan. 1921 BO, L Beguin 1894 Galela 10 2 Dec. 1921 BO, L Beguin 2037 Sao Tobaroe 60 19 June 1922 BO, L, U Pandanus polycephalus Bangun 520 Tjetju, Weda Bay 17 15 Jan. 2013 BO Bangun 875 Lelilef. Weda Bay 107 14 Sept. 2013 BO, G, L, MO Beguin 1792 Galela 1 28 Sept. 1921 L Beguin 1966 Galela 20 19 May 1922 BO, L Beguin s.n. Galela, Sao Tobaroe 1920 BO Merello 3304 Weda Bay 33 10 Oct. 2012 BO, MO Merello 3312 Forest along Gomdi river 20 11 Oct. 2012 BO, G, L, MO Taylor 424 Jikolamo 6 June 1981 SAN Teijsmann 87 s.l. s.d. L Vriese 18 s.l. s.d. L Yoshida 2390 From Limau to Soa, Galela Dist. 12 Oct. 1976 L Pandanus tectorius Merello 3311 Gwomdi , Weda Bay 17 11 Oct. 2012 BO, G, K, L, MO Candollea 70, 2015 The genus Pandanus (Pandanaceae) on Halmahera Island (Moluccas, Indonesia)– 187 sub-populations are discovered, the estimated AOO and EOO for the species would increase, and reassessment of the conservation status of the species may result in a lower category of threat. Furthermore the establishment of conservation offset sites to mitigate mining and associated impacts that encompass natural subpopulations of the species would decrease the level of threat to the species. Notes. – Pandanus beguinii can easily be distinguished by its small, slender habit, its small, terminal, erect syncarp of truncate drupes topped by a dark turret with a flat to concave apex, bearing a lateral minute bicornuate stigma, and its singlespiked staminate inflorescence (Fig. 3A-C, 5). The new species is a member of the morphologically well-defined Pandanus sect. Maysops H. St. John, that is characterized by “cephalia resembling an ear of maize, with flat-topped drupes” (Jebb, 1992: 116). This section, distributed throughout the Malesian region from the Moluccas to Papua New Guinea, is also present in the Bismarck Archipelago, the Solomon Islands and northern Queensland in Australian (Stone, 1992). Within this section, P. beguinii is similar to P. kosteri B. C. Stone in being miniature in all parts compared to all other species of the section, which develop into relatively robust trees. Pandanus beguinii can nevertheless be easily distinguished by its broader leaves, mostly 5-8cm wide across the middle (vs.2.5-3.7cm in P. kosteri), its larger syncarp (10-15 × 5-6cm vs. 6.5 × 3.5cm), its simple staminate inflorescence with yellow bracts (vs. a compound inflorescence comprised of 6-7 spikes with red bracts) and its staminate flowers with their stamens shortly basally connate (vs. fused, i.e. forming a phalange) (see Stone, 1987: 436, Fig. 5). Paratypi. – indoneSia. Prov. North Maluku [Halmahera Isl.] : Loleo Poeloe [Loypoloy] Rao, 13.X.1921, Beguin 1819 (BO) ; ibid. loc., 30 m, 14.X.1921, Beguin 1822 (BO) ; ibid. loc., 40 m, 15.X.1921, Beguin 1825 (BO) ; Soa Tobaroe, 60 m, 24.VI.1922, Beguin 2044 (BO) ; ibid. loc., 60 m, 28.VI.1922, Beguin 2067 (BO) ; ibid. loc., 29.VI.1922, Beguin 2069 (BO, L [L1195381]) ; ibid. loc., 29.VI.1922, Beguin 2071 (BO) ; ibid. loc., 60 m, 29.VI.1922, Beguin 2071 (BO) ; ibid. loc., 29.VI.1922, Beguin 2107 (L [L1195381]) ; ibid. loc., 6.VIII.1922, Beguin, 2167 (BO) ; Central Halmahera, Weda Bay, on the road to Bukit Limber, 00°31’01”N 127°59’41”E, 600 m, 15.IV.2013, Bidault et al. 1089 (BO, G, L, MO) ; Weda Bay, Sagea Lagoon, 00°29’35”N 128°04’16”E, 29.I.2013, 15 m, Callmander et al. 1086 (BO, G, L, MO-6465090) ; East Halmahera, Weda Bay, Pinto, 00°38’10”N 128°09’05”E, 340 m, 1.II.2013, Callmander et al. 1090 (BO, G) ; East Halmahera, Weda Bay, Kao Rahai, 00°40’22”N 127°58’31”E, 366 m, 9.XII.2012, Merello et al. 3539A (BO, L, MO-6444045, MO-6444047). Pandanus benstoneoides Callm., Buerki & Phillipson, spec. nova (Fig. 2A-D, 4B, 6, 7). Typus: indoneSia. Prov. North Maluku [Halmahera Isl.] : East Halmahera, Weda Bay, Pinto, 00°38’10”N 128°09’05”E, 340 m, 1.II.2013, Callmander et al. 1096 (holo- : G [G00341588] ! ; iso- : BO!, BM!, L!, MO-6465097!, MO-6465098!). Haec species a congeneris inflorescentiis lateralibus axillaribus, staminibus liberis atque drupis unilocularibus stigma filiforme adpressum rectum undulatumve syncarpi apicem versus dirigens gerentibus distinguitur. Treelet to 4-8(-15) m tall, stem c. 15cm in diam., with a large cone of few proproots at base, up to 2 m high. Leaves linear-attenuate, mostly (110-)130-150(-160)cm long, 4.5-5.5cm wide in the middle, apex gradually attenuate, ending with a flagellum (c. 7cm long) ; sub-coriaceous, glaucous abaxially; longitudinal veins visible on both surfaces; marginal prickles borne 4-9cm above base to apex, black near sheath, antrorse, erect, 0.8-1.5mm long in the lower third, 0.5-2mm apart, 0.5-1mm long in the mid-third, 1-3mm apart, to 0.2-0.5mm long in the distal third, 0.2-1mm apart, appressed in the distal third; midrib armed from 35-50cm from base, prickles > 1mm long towards the apex, irregularly spaced but mostly 2-4mm apart in the middle and 1-2mm towards the apex; prickles generally absent on apical ventral pleats, sometimes 5-10 minute prickles towards the apex; sheath straight, (4-)5-7(-8)cm long, 4.5-6cm wide at apex, 5-8cm at base. Infructescence lateral, comprising a solitary patent syncarp born in the axil of the leaves, several present in each fertile leaf rosette; each syncarp ovoid, obtuse at apex, 3-angled, 7-10cm long, 5-7cm wide, peduncle short, flattened dorso-ventrally, c. 7cm long, 1-1.5cm thick at apex, bearing multiple sub-coriaceous yellow bracts, broadly naviculate, the proximal ones c. 11cm long, the distal one 7cm long, crowded at peduncle apex, covering the syncarp. Drupes numerous, 500-800, 13-15mm long, 3-4mm wide, 4-5mm thick, 1-celled, 5-6-angled, linear, pileus short, 1-2mm high, with a flat to slightly concave apex with a rugose surface, bearing an appressed, straight to wavy, filiform stigma on its upper edge; stigma towards the syncarp apex, (10-)15-20(-30)mm long, 1mm wide at base, 0.5mm at apex; stigmatic groove ventral (adaxial), running along the proximal two thirds of the stigma. Endocarp bony, median, c. 4mm long, obovate, lateral walls >0.4mm thick, extending laterally toward the apex of the drupe; proximal mesocarp c. 3mm long, fibrous, distal mesocarp c. 3-5mm long, chambered, seed locule c. 1.5×3mm, obovate to ovate. Staminate inflorescence c. 18cm long (immature) ; with a single terminal spike on a peduncle c. 7cm long, covered with narrowly navicular yellow bracts, proximal ones c. 14 × 3cm; distal ones 8 × 2cm; spike yellow, c. 7 × 1.5cm, linear, composed of crowded and congested flowers, composed of free stamens; stamens c. 5.5mm long; filament, 4 × 0.5-0.7mm, oblong, often larger at apex (c. 1mm wide) ; anther c. 1.5 × 1mm, sometimes apiculate with a long slender apiculus c. 1mm long. Etymology. – This new species is named after its resemblance to species of the genus Benstonea, a genus characterized 194 –The genus Pandanus (Pandanaceae) on Halmahera Island (Moluccas, Indonesia) Candollea 70, 2015 Acknowledgments The authors are grateful to Joeni Setijo Rahaju and Deby Arifiani at the Herbarium Bogoriense (BO) for their support and assistance, and to PT Weda Bay Nickel for the opportunity to participate in their Flora Inventory Programme on Halmahera, with special thanks to Tjut Fatisa Bangun, Bahar Fabanyo, Iska Gushilman, Idris Haris, Samsul Lasut, Gavin Lee, Roji Mahroji, Edi Permana, and Bilal Sau. We want to sincerely thank Ehoarn Bidault, SusanaArias Guerrero, Pete Lowry, Mary Merello and Ed de Vogel for their interest, tireless support and help in this study. We also thank Roy Gereau for preparing the Latin diagnoses. We are finally grateful for the valuable comments on the original manuscript provided by Henk Beentje. References Backer, C. A. (1936). 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