PALM S Baker & Heatubun: and Supiori Palms Vol. 56(3) 2012

WILLIAM J. B AKER New Palms Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom from Biak [email protected] and AND

CHARLIE D. H EATUBUN Supiori, Fakultas Kehutanan, Universitas , Jl. Gunung Salju, Amban, Western 98314, Papua Barat, [email protected]

1. View from Samber across the limestone cliffs of western Biak. (Photo: W.J. Baker)

The Indonesian of Biak and Supiori sit at the mouth of , the broad arc of ocean to the north of the neck of New Guinea’s Bird’s Head Peninsula. Already known for their remarkable animal endemism, the islands are home to four unique palm species, three of which are described here for first time.

PALMS 56(3): 131 –150 131 PALM S Baker & Heatubun: Biak and Supiori Palms Vol. 56(3) 2012

Of the many islands scattered around already been introduced to cultivation prior Cenderawasih Bay (also known as Geelvink to its formal description but was poorly known Bay), Biak, Supiori, and stand in the wild until these expeditions. The out on account of their relatively large size. remaining three new species are described here Yapen, a land bridge , sits close to the for the first time. New Guinea mainland to which it was once As a whole, the palm flora of Biak is not connected, while the remaining three are particularly rich (Table 1). Nineteen species oceanic in origin and located in the north of have been recorded, with all non-endemic the bay. Biak forms an island pair with the species being shared with mainland New smaller Supiori, the two being separated only Guinea. Widespread New Guinean species are by a narrow channel, and Numfor is located present, such as Arenga microcarpa , Caryota some 60 km to the south-west. Together, Biak rumphiana and Hydriastele costata , as well as and Supiori measure some 125 km in length two that are distinctive western New Guinean and 40 km at the widest point. In common species, Pigafetta filaris and Pinanga rumphiana with other islands in the region, they are (Figs. 2 & 3). The Biak palm flora also includes dominated by rugged limestone (Fig. 1) several widespread New Guinea rattan species perforated by sink-holes and caves, with such as Calamus aruensis , C. heteracanthus , C. sporadic occurrences of other geologies. Biak pachypus , C. vitiensis , C. zebrinus and Korthalsia played an important role in World War II and zippelii . Of Biak’s non-endemic palms, only was the scene of some of the bloodiest fighting Calamus dasyacanthus and Hydriastele brassii in the Pacific. The Japanese made use of some (Back Cover) could be regarded as less well of Biak’s caves as bases and defensive position, known. Our knowledge of the palm flora of but were ultimately defeated by Allied forces Supiori is less complete. We expect that it is after a bitter stand-off. Biak’s large airstrip dates likely to be broadly similar to that of Biak, back to that time and has allowed Biak to although the presence of a high peak (>1000 function as a hub for air travel in the region. m) and somewhat different geology (Masria et al. 1981) may result in the occurrence of The relative accessibility of Biak by boat and additional montane species. air has no doubt contributed to the degradation of the island’s rain forest, much In contrast to the relatively ordinary non- of which has now been destroyed. The North endemic palm flora, the endemic palms are Biak Island Reserve encompasses a portion of remarkable in various ways. Hydriastele the northern tip of the island and much of dransfieldii was originally described in Supiori is also protected, although the Siphokentia (Baker et al. 2000), a genus that was later reduced into synonymy with effectiveness of these reserves is not clear. Some Hydriastele , following in depth molecular unprotected areas of the interior of the island phylogenetic studies (Baker & Loo 2004, Loo also appear to retain good forest, though et al. 2006). While this taxonomic change is further analysis of the condition of the corroborated by morphology, the two species remaining primary vegetation is urgently formerly accepted in Siphokentia (H. dransfieldii required. and the Moluccan H. beguinii ) remain a The plants of Biak and Supiori are poorly distinctive pair of sister species within known. The palms, however, have been the Hydriastele , characterized by the fused petals subject of a series of expeditions to Biak, two and sepals in the female flower. We found H. led by Rudi Maturbongs with colleagues from dransfieldii to be widespread on limestone in Universitas Negeri Papua in 1998 and 2001, Biak and abundant in places. Occasionally, it and three involving the authors and various can even be observed in disturbed roadside colleagues from Universitas Negeri Papua, vegetation, although it is unlikely to be Herbarium Bogoriense and the Royal Botanic regenerating in such secondary habitats. Gardens, Kew in 2000, 2009 and 2010. The Hydriastele dransfieldii has been reported (sight 2001 expedition also included a short visit to records only) from Numfor and Supiori, but Supiori. As a result of these expeditions, we the species is not known from Yapen or have obtained sufficient material for the mainland New Guinea. description of four remarkable new endemic The three remaining endemic species, species, three from Biak and one from Supiori. described as new to science below, are The first of these, Hydriastele dransfieldii was apparently far less common on Biak and face described following the expeditions led by a greater threat of extinction. Like H. Maturbongs (Baker et al. 2000). This palm had dransfieldii , they are palms of limestone, two

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133 PALM S Baker & Heatubun: Biak and Supiori Palms Vol. 56(3) 2012

2. Pigafetta filaris , a common palm in disturbed vegetation on Biak. (Photo: W.J. Baker) of which (like H. dransfieldii ) show bio- Biak Island: forest on the road side, main road geographic relationships with species on from North Biak Nature Reserve to Biak town, islands to the north-west, remarkable in view July 2009, Heatubun et al. 971 (holotype K, of the much greater species pool in nearby isotypes BO, FTG, MAN, NY). (Figs. 4–11) New Guinea to the south. All three represent significant additions to our knowledge of the Diagnostic characters: Adonidia maturbongsii palms of the New Guinea region and serve to is distinguished from A. merrillii by the arching focus attention on the importance and plight leaf, broad, pendulous leaflets in a single plane, of Biak’s dwindling rain forest. wide, concave leaflet tips and staminate flowers containing 30–32 stamens. Taxonomic treatment Medium, solitary, mid-story to emergent palm. 1. Adonidia maturbongsii W.J.Baker & Stem 10–15 m tall, 10–20 cm in diam., Heatubun, sp. nov. Type: Indonesia, Papua, tapering towards apex, surface brown with

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3. Pinanga rumphiana , Biak. (Photo: W.J. Baker) white blotches, leaf scars prominent, 26–45 cm long, channelled adaxially, rachis internodes 2–4 cm apart. Leaves ca. 10 in 2.5–3 m long, indumentum as on sheath; crown, arching; sheath 60–70 cm long, pale, leaflets 25–30 pairs each side of the rachis, dull green, with thin grey scurfy indumentum regularly arranged (or somewhat subregularly), with scattered purple-brown scales, somewhat in one plane, drooping or pendulous in eroded or fibrous at mouth, forming emergent individuals, slightly discolorous, crownshaft 80–90 cm × 10–12 cm; petiole with persistent reins attached to lowermost

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4. Adonidia maturbongsii . A. Habit. B. Leaf apex. C. Mid-leaf portion. D. Inflorescence. E. Portion of rachilla with triad. F. Staminate flower. G. Pistillate flower in section. H. Fruit whole and in section. I. Endocarp showing longitudinal fibers. Scale bar: A = 2 m; B–D = 6 cm; E = 1.5 cm; F, G = 7mm; H, I = 7 mm. From Heatubun et al. 971 , except A from photograph and F from Baker et al. 1338 . Drawn by Lucy T. Smith.

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5. Adonidia maturbongsii on the outskirts of Kota Biak. (Photo: W.J. Baker) pair of leaflets, with minute brown punctate Inflorescence 60–70 cm long, infrafoliar, scales and scattered medifixed ramenta protandrous, divaricate, patent, deflexed in abaxially; middle leaflets 40–49 cm long, 9–12 fruit, branched to 4 orders, axes white, rubbery, cm wide, oblanceolate, cucullate, apex with caducous floccose orange-brown obliquely praemorse, concave, transverse indumentum when young; prophyll 24–26 cm veinlets inconspicuous; terminal leaflets linear long, 6–8 cm wide, greenish white, splitting or narrowly elliptic, apex truncate, praemorse. apically, caducous later; first peduncular bract

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31–35 cm × 5–7 cm, attached 15–20 mm above black-brown scales basally; primary branches prophyll insertion, exserted from prophyll 25–28, longest primary branch (basalmost) apex and enclosing inflorescence prior to 40–65 cm; rachillae 8–19 cm long, 1.5–3.5 mm anthesis, caducous later; peduncle 8–14 cm in diam., triads 3–9 mm apart, spirally long, 2–2.5 cm wide, scurfy indumentum of arranged. Staminate flower 6.5–8 mm long,

6 (top). Adonidia maturbongsii , a young specimen emerging from the canopy with less pendulous leaflets than adults, Samber forest, Biak. 7 (bottom). Adonidia maturbongsii leaf. (Both photos: W.J. Baker)

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2.5–3.2 mm in diam. in bud; sepals 2–2.4 mm long; filaments 1.5–4 mm long, briefly connate long, ca. 3 mm wide, rounded, thickened; at base, awl-shaped; anthers 3–3.8 mm long, petals 7–7.5 mm long, ca. 3 mm wide, bony, 0.5–0.8 mm wide, dorsifixed near the base, narrowly elliptic; stamens 30–32, 4.5–6 mm dehiscence latrorse, connective dark; pistillode

Adonidia maturbongsii. 8 (upper left). Inflorescence at staminate anthesis, held by Charlie Heatubun. 9 (upper right). Staminate flowers. 10 (bottom). Infructescence. (All photos: W.J. Baker)

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11. Adonidia maturbongsii , fruits. (Photo: W.J. Baker) ca. 5 mm long, 1.2–1.4 mm in diam., ca. 4.5 mm in diam., borne in proximal half lageniform. Pistillate flower 4.5–5 mm long, to two-thirds of the rachilla; sepals 3–4 mm

140 PALM S Baker & Heatubun: Biak and Supiori Palms Vol. 56(3) 2012 long, 4–4.5 mm wide, thickened, rounded; northern Borneo (Zona et al. 2011). Although petals 4–4.5 mm long, 3–3.5 mm wide, similar the study was based on only two DNA regions to sepals; staminodes few, minute, tooth-like; and the relationships only moderately gynoecium ca. 4 mm long, ca. 3 mm in diam., supported by the data, we describe the new pyriform, stigmas at anthesis not seen. Fruit species as Adonidia maturbongsii as the best 24–31 mm long, 14–16 mm in diam., ellipsoid, solution given the available data and because ripening through orange to red, perianth of morphological similarities discussed below. cupule clasping; endocarp 23–30 mm long, Generic limits in subtribe Ptychospermatinae 12–12.5 mm in diam., straw-colored with thick are fine and sometimes problematic (Zona longitudinal fibers, closely adhering to seed. 1999, Dransfield et al. 2008), as evidenced by Seed 14–20 mm long, 9.5–12 mm in diam., the initial, but erroneous assignment of this ellipsoid; endosperm ruminate; embryo basal. new species to Drymophloeus , a genus that has Distribution: Scattered throughout Biak experienced substantial changes in Island. circumscription recently (Zona et al. 2011). Nevertheless, A. maturbongsii and A. merrillii Habitat: Lowland forest on limestone with share a combination of features that lends thin soils and many sink holes, 80–170 m support to a close relationship between the elevation. two. Both species are moderately robust palms Vernacular names: Manjek (Biak dialect) of limestone habitats that bear white inflorescences branched up to four orders. Uses: Stem used for flooring and pillars in They produce red fruit with endocarps covered traditional houses. in pale, flattened, longitudinal fibers Conservation status: Endangered (EN B1, 2 (a, interspersed with finer fibers and seeds with b [i, ii, iii, iv, v]), C2a (i); IUCN 2001). Though ruminate endosperm. Nevertheless, A. widespread on Biak, many sites for this species maturbongsii is very different from its congener, are threatened by ongoing forest degradation. most obviously in its arching leaf with broad, The species is protected in the North Biak pendulous leaflets in a single plane with wide, Nature Reserve. concave, praemorse tips (in contrast to the ascending, narrower leaflets in slightly Specimens examined: Indonesia, Papua, Biak different planes with less conspicuously Island: forest on the road side, main road from praemorse tips in A. merrillii ). In addition, the North Biak Nature Reserve to Biak town, July staminate flowers of A. maturbongsii contain 2009, Heatubun et al. 971 (holotype K, isotypes 30–32 stamens compared to 45–50 in A. BO, FTG, MAN, NY); North Biak Nature merrillii . Reserve, Sansundi village, September 1998, Maturbongs et al. 559 (BO, K, MAN), Maturbongs Adonidia merrillii is a geographically disjunct et al. 560 (BO, K, MAN); Samber forest, July species, occurring to the west of Wallace’s Line 2009, Baker et al. 1336 (BO, K, FTG, MAN), whereas all other Ptychospermatinae occur to Baker et al. 1338 (BO, K, FTG, MAN); locality the east of this important biogeographic uncertain (given incorrectly on the label as interface (Baker & Couvreur 2012). The Merauke district, but number sequence expansion of the genus elaborates this indicates the collector was active in Biak), June biogeographic story. The link between New 2001, Maturbongs et al. 686 (AAU, BO, FTG, K, Guinea and the Philippines has been explained MAN). by westward stepping-stone dispersal along the Philippine-Halmahera arc during the Notes: It has been known for some time that Neogene (Zona et al. 2011), which may also an undescribed species from subtribe account for similar biogeographic links in Ptychospermatinae occurs on Biak (Baker pers. other taxa, such as the palm genera obs. 2000, Zona 2000). The species appears to Heterospathe and Orania , and Sararanga in the have been first recorded by Greg Hambali who Pandanaceae (Baker et al. 1998, Norup et al. introduced it to cultivation as Drymophloeus 2006). “veitchioides,” an unpublished name under which it persists in some collections today. Adonidia maturbongsii is named for our friend However, new molecular phylogenetic data and collaborator Rudi Maturbongs of provide evidence that the species is most Universitas Negeri Papua, the collector of the closely related to Adonidia merrillii , the sole first herbarium specimens of the new species, species of a hitherto monotypic genus in recognition of his contributions to palm restricted to parts of the Philippines and far exploration in Biak.

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2. Heterospathe porcata W.J. Baker & only, bracteole forming conspicuous perianth- Heatubun, sp. nov. Type: Indonesia, Papua, like cupule; sepals 3, ca. 2.5 mm long, ca. 3 : North Supiori Nature Reserve, mm wide, concave, imbricate; petals 3, ca. 3.6 Fanjur village, June 2001, Maturbongs et al. 680 mm long, ca. 2.5 mm wide, concave with (holotype K; isotypes AAU, BO, CANB, LAE, short, triangular apical lobe, imbricate; MAN). (Fig. 12) staminodes ca. 2, linear, ca. 0.6 mm long; gynoecium ca. 3.5 mm long, ca. 1.5 mm in Diagnostic characters: Heterospathe porcata is diam., ellipsoid, stigma inconspicuous. Fruit distinguished from other species in the genus ca. 2 cm long, ca. 9 mm diam. (measured from by its inflorescence with elongate peduncle, dry material), ellipsoid, pericarp shrinking the peduncular bract inserted in the proximal around endocarp ridges when dried, stigmatic half of the peduncle, and the fruit with a bony remains eccentrically apical, red; perianth endocarp with 6–7 longitudinal ridges. cupule clasping; endocarp ca. 2 cm long, ca. Slender, ?solitary, understory palm. Stem to 8.5 mm in diam., ellipsoid, dark brown, bony, ca. 6 m tall, ca. 3 cm in diam.; leaf scars with 6–7 thickened, fibrous ribs radiating from prominent; internodes 1–2 cm. Leaves 16 in the stigmatic remains and running the length crown; sheath open, margins eroded, not of the fruit to the base. Seed immature, star- forming crownshaft; petiole ca. 50 cm long, shaped in section, conforming to inner adaxially channeled; rachis ca. 100 cm long, contours of endocarp; endosperm ruminate; with sparse, floccose indumentum of orange- embryo basal. brown scales throughout; leaflets ca. 40 each Distribution: Recorded from only one locality side of rachis, regularly arranged, borne up to on Supiori Island. 3.5 cm apart, somewhat discolorous, basifixed ramenta scattered on proximal part of adaxial Habitat: Secondary limestone forest surface of midribs and major veins; middle dominated by Myristica , Intsia and Lepiniopsis leaflet ca. 45 cm long, 2–2.3 cm wide, linear, at an elevation of ca. 30 m. somewhat sigmoid at tip and base, tip Vernacular names: Not known. narrowly attenuating, transverse veinlets inconspicuous. Inflorescence 124–138 cm Uses: The stem is used for making bows and long, interfoliar, elongate, brush-like, branched the fruit is used as a betel nut substitute. to 2–3 orders; prophyll ca. 39 cm long, 1.5–2 Conservation status: Data deficient (IUCN cm wide, splitting apically, with lepidote 2001). Current knowledge of populations of indumentum of dark scales; first peduncular this palm and the condition of the forests on bract, ca. 69 cm long, ca. 2 cm wide, similar Supiori is insufficient for a conservation to prophyll, splitting apically to one side, assessment to be completed at this time. indumentum as prophyll, attached one third to halfway above the peduncle base; peduncle Specimens examined: Indonesia, Papua, 102–108 cm long, 5.5–7.5 mm wide, with thin, Supiori Island: North Supiori Nature Reserve, dark brown tomentum and throughout all Fanjur village, June 2001, Maturbongs et al. 680 inflorescence branches; primary branches (holotype K; isotypes AAU, BO, CANB, LAE, 9–13, to 28 cm long, 1.5–4 cm apart, with up MAN). to 14 rachillae; rachillae 10–17 mm long, Notes: Heterospathe has two centers of diversity, 0.9–1.5 mm in diam., swept forward at an the Philippines and New Guinea. In New acute angle to the rachis, sinuous, with thin Guinea, species diversity is biased towards the dark brown tomentum; rachilla bracts eastern half of the island and montane inconspicuous; triads 2–3.5 mm apart, spirally elevations – around 90% of all New Guinea arranged. Male flower ca. 2 mm long, ca. 1.8 Heterospathe specimens have been collected mm in diam. in young bud; sepals 3, ca. 1 mm from and more than 75% long, ca. 1.5 mm wide, concave, imbricate; from elevations above 500 m. An undescribed petals 3, ca. 1.5 mm long, ca. 1.3 mm wide, species from the lowlands of an offshore island cucullate, valvate; stamens 6; filaments ca. 0.6 of is thus an unexpected mm long, connate in a ring at base, linear; discovery. anthers ca. 0.8 mm long, ca. 0.3 mm wide, oblong, dorsifixed, connective dark; pistillode Superficially, H. porcata is similar to 0.6 mm long, 0.3 mm in diam., cylindrical. Heterospathe elegans , although the nearest Female flower ca. 4 mm long, ca. 3.5 mm in record for this species is some 500 km away to diam., borne in proximal half of the rachillae the south-east (Trudgen & Baker 2008). The

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12. Heterospathe porcata . A. Leaf apex and mid-leaf portion. B. Inflorescence in two halves. C. Detail of rachilla. D. Rachilla with pistillate flowers, one removed to show large floral bracteole. E. Staminate flower whole and in section. F. Pistillate flower in section. G. Endocarp in two views. Scale bar: A, B = 8 cm; C, D = 7 mm; E = 2 mm; F = 3mm; G = 1 cm. From Maturbongs et al. 680 . Drawn by Lucy T. Smith.

143 PALM S Baker & Heatubun: Biak and Supiori Palms Vol. 56(3) 2012 two species share a slender habit, finely structures. The combination of its geographical pinnate leaf and inflorescence with elongate and elevational distribution, and almost peduncle and branches clustered near the unique reproductive morphology renders H. apex. Heterospathe porcata appears to be a taller porcata a very surprising novelty indeed. palm, recorded as 6 m in contrast to the 3. Hydriastele biakensis W.J.Baker & reported maximum of 2.5 m for H. elegans . Heatubun sp. nov. Type: Indonesia, Papua, The inflorescence is also quite different, being Biak Island: Oridek district, Wadibu village, branched to 2 or 3 orders (1 or 2 orders in H. July 2009, Baker et al. 1342 (holotype K, elegans ), the rachillae being finely sinuous isotypes AAU, BO, MAN). (Figs. 13–19) (more irregularly so in H. elegans ) and the first peduncular bract being rather long and Diagnostic characters: Hydriastele biakensis is inserted one third to halfway along the distinguished from other species in the genus peduncle from the base (shorter in H. elegans by its large size, the recurving leaves with and inserted in the distal quarter of the ascending leaflets and acute or notched leaflet peduncle). Unusually, the fruit contains a thin, apices, the often somewhat distorted prophyll bony endocarp with 6 or 7 thickened, parallel, bearing pronounced pithy keels, the peduncle fibrous ridges running the full length of the abruptly constricting at the prophyll scar, the fruit (hence the species epithet porcata , inflorescence branched to four orders, the meaning ridged). These reveal themselves highly sinuous rachillae, and the congenitally when the ellipsoid fruit, which are larger than open staminate flowers. the globose fruit of H. elegans , are dried and the Robust, solitary, canopy palm. Stem ca. 15 m mesocarp shrinks around the ridges. The space tall, ca. 30 cm in diam., leaf scars prominent, with the endocarp is somewhat star-shaped in internodes 3–7 cm, surface brown. Leaves ca. cross section and, consequently, so is the seed, 18–24 in crown, strongly recurved; sheath ca. in contrast to the globose seed of H. elegans . 170 cm long, pale green with white waxy Within Heterospathe, the only species with a indumentum, striate near mouth, forming comparable endocarp is H. longipes of Fiji, crownshaft 200–270 cm long, 26–27 cm wide; though this has much more extremely ornate rachis 280–300 cm long, petiole 47–50 cm

13. Hydriastele biakensis , cultivated at Marau, Biak. (Photo: W.J. Baker)

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14. Hydriastele biakensis , cultivated at Marau, Biak. (Photo: W.J. Baker) long, 3.5–5 cm wide, channeled adaxially, minute dark dots throughout; leaflets ca. 65 petiole and lower rachis yellowish green, each side of rachis, regularly arranged, strongly petiole and rachis bearing scattered to dense ascending, concolorous, with minute dark dots brown, caducous, felty indumentum and abaxially, brown, basifixed ramenta attached

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15. Hydriastele biakensis . A. Leaf apex and mid-leaf portion. B. Inflorescence base. C. Inflorescence first order branch. D. Rachilla showing congenitally open flowers. E. Rachilla with pistillate flowers at anthesis. F. Staminate flower. G. Pistillate flower whole and in section. H. Sinuous rachilla with fruit attached. I. Fruit in section. Scale bar: A,C = 6 cm; B = 8 cm; D, H = 1.5 cm; E, I = 1 cm; F = 5 mm; G = 3 mm. A–C from Baker et al. 1342, D–I from Heatubun et al. 970. Drawn by Lucy T. Smith.

146 PALM S Baker & Heatubun: Biak and Supiori Palms Vol. 56(3) 2012 to basal, abaxial portion of midrib; middle mm wide, narrowly triangular, variously leaflets 121–126 cm long, 3–4.5 cm wide, twisted and sinuous, briefly adnate to linear, transverse veinlets conspicuous, apices receptacle, white; stamens 6, 4.5–5.5 mm long, narrowly acute; terminal segments linear, with white; filaments ca. 0.5 mm long, narrowly apices notched, not praemorse. Inflorescence conoid; anthers 4–5 mm long, 0.8–1.2 mm 95–100 cm long, infrafoliar, ?protandrous, wide, oblong to sinuous, basifixed, dehiscence horsetail-shaped, erect, branched to 4 orders, latrorse; pistillode minute, pyriform. Pistillate axes white on emergence, turning green; flower ca. 2.5 mm long, 2.5–2.8 mm in diam., prophyll 70–107 cm long, 15–18 cm wide, borne throughout the rachillae; sepals green, often somewhat sinuous, appearing imbricate, ca. 1 mm long, 2.5–2.8 mm wide, distorted, keels pithy, with thin, white rounded, white; petals 2–2.5 mm long, 2.5–3 indumentum; first peduncular bract, ca. 70 cm mm wide, strongly imbricate, rounded, white; long, ca. 5.5 cm wide, similar to prophyll, attached 2.5–4.5 cm above prophyll insertion; staminodes 3, minute, paddle-shaped; peduncle 10–21 cm long, 8–8.5 cm wide at gynoecium ca. 2 mm long, ca. 1.5 mm in base, narrowing sharply above prophyll diam., globose; stigma minutely trifid. Fruit insertion to 3–3.5 cm, prophyll scar 9.5–12 mm long, 5–6 mm in diam., oblong- conspicuous with rounded “shoulders”; ellipsoid, red, perianth cupule clasping, primary branches 20–22, the longest endocarp thin, tough, closely adhering to seed. (basalmost) to 70 cm; rachillae 32–48 mm Seed 7.5–8.2 mm long, 4–4.3 mm in diam., long, 2–3 mm in diam., sinuous, especially cylindrical; endosperm homogeneous; embryo distally, triads 2–3 mm apart, decussate. basal. Staminate flower 5–6.5 mm long, 2.5–4 mm in diam. in bud, variously flattened and Distribution: Known from few specimen distorted, congenitally open; sepals connate localities and sight records on the south and in a shallow cup with three triangular lobes ca. western coast of Biak and Auki Island on the 0.5 mm long, white; petals 5–6 mm long, 1–1.5 nearby Padaido Islands.

Hydriastele biakensis . 16 (left). Inflorescences. 17 (right). Infructescences. (Both photos: W.J. Baker)

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Habitat: Coastal forest on limestone near to sea level, sometimes on limestone cliffs close to the beach. Vernacular names: Arwaf (Biak dialect) Uses: The stem is used for flooring and the leaf sheaths for making baskets. Conservation status: Endangered (EN B1, 2 (a, b [i, ii, iii, iv, v]), C2a (i); IUCN 2001). This species is known only from coastal limestone forest which is severely degraded on Biak, especially on the south coast. Larger populations have been observed on the west coast and Padaido Islands. Specimens examined: Indonesia, Papua, Biak Island: Oridek district, Wadibu village, July 2009, Baker et al. 1342 (holotype K, isotypes AAU, BO, MAN), Heatubun et al. 970 (BO, K, MAN, NY). A sterile specimen may also represent this species: northern Biak, Wari village, September 1998, Maturbongs et al. 574 (BO, K, MAN). Notes: We became aware of this beautiful species during the brief visit to Biak in 2000 (Baker pers. obs.) when it was seen cultivated near the now ruined Marau Beach Hotel and persisting wild as a few scattered individuals in cleared areas near the south coast. At that time, 18. Hydriastele biakensis , congenitally open male it was regarded as a species of Gulubia (now a flowers prior to anthesis. (Photo: W.J. Baker) synonym of Hydriastele ) and suspected to be undescribed when compared with the species This character, alongside fruit structure, was treated in Essig’s (1982) monograph of the considered diagnostic for the genus Gulubiopsis genus. Unable to collect material at that time, in which H. palauensis was originally described we had to wait until 2009 for an opportunity (Beccari 1924, Beccari & Pichi-Sermolli 1955). to make complete specimens for herbarium Moore and Fosberg (1956) deemed these and laboratory study. features inadequate to justify generic status, reducing Gulubiopsis into synonymy with To determine the relationships of the new Gulubia , which was later sunk into Hydriastele species, we exploited an earlier phylogenetic (Baker & Loo 2004). study of Hydriastele (Loo et al. 2006). Following the protocols of Loo et al., we generated new In addition to morphological similarities, the DNA sequence data for H. biakensis of the two two species occupy similar coastal limestone low-copy nuclear genes PRK and RPB2, habitats. Moreover, Biak is among the closest integrated these new data within their of the Malesian islands to , although published dataset and repeated their analyses. almost 1000 km of clear ocean exists between Hydriastele biakensis was strongly supported as the two. Nevertheless, the two are clearly sister species of H. palauensis . There are distinct species. Hydriastele biakensis is much morphological similarities between the two more robust than H. palauensis, for example species in general appearance, such as the with stem diameter, leaf length, leaflet number, strongly recurved leaves, ascending leaflets leaflet length, sheath length and inflorescence with acute or notched (but not conspicuously being twice the size or more in the former than praemorse) apices and the glaucous that reported for the latter (Moore & Fosberg crownshaft. The two also share the unusual 1956, Essig 1982). The contrast is most clear feature of the staminate flowers being in the inflorescence which, as well as being congenitally open in bud due to the large size much smaller in H. palauensis , lacks the of the stamens relative to the petals (Fig. 18). striking “shoulders” formed by the abrupt

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19. Hydriastele biakensis , on low limestone cliffs above the beach at Wadibu, Biak. (Photo: W.J. Baker)

149 PALM S Baker & Heatubun: Biak and Supiori Palms Vol. 56(3) 2012 constriction of the peduncle at the prophyll DRANSFIELD , J., N.W. U HL , C.B. A SMUSSEN , W.J. scar, and the highly sinuous rachillae (see p. BAKER , M.M. H ARLEY AND C.E. L EWIS . 2008. 107), and in the material available to us Genera Palmarum – the Evolution and (Lorence et al. 8304 [PTBG]) is branched to two Classification of Palms. Royal Botanic rather than four orders. Gardens, Kew, Richmond.

Acknowledgments ESSIG , F.B. 1982. A synopsis of the genus We are grateful to many colleagues who Gulubia . Principes 26: 159–173. provided comments on the manuscript, IUCN. 2001. IUCN Red List Categories and additional information or support in the field, Criteria: Version 3.1. Species Survival lab or herbarium: Theo Ampnir, Deby Arifiani, Commission, IUCN, Gland, Switzerland and Steve Bachman, Jim Clarkson, John Dransfield, Cambridge. Rachel Engstrand, Lauren Gardiner, Jeff Marcus, Justin Moat, Himmah Rustiami, Tim LOO , A.H.B., J. D RANSFIELD , M.W. C HASE AND W.J. Utteridge and Scott Zona. We are especially BAKER . 2006. Low-copy nuclear DNA, grateful to Rudi Maturbongs for the important phylogeny and the evolution of dichogamy contributions he made in the earlier in the betel nut palms and their relatives explorations of Biak’s palms. Fieldwork was (Arecinae; Arecaceae). Molecular Phylo- funded by the Tobu fund, the BAT Biodiversity genetics and Evolution 39: 598–618. Partnership and the Royal Botanic Gardens, MASRIA , M., N. R ATMAN AND K. S UWITODIRDJO . Kew. This paper is dedicated to the late Charles 1981. Geologi lembar Biak, Irian Jaya (The H. Uhl (1918–2010), a veteran of the war in geology of the Biak Quadrangle, Irian Jaya). Biak and husband of the great palm biologist, Geological Research and Development Natalie W. Uhl. Centre, Bandung. References MOORE , H.E. AND F.R. F OSBERG . 1956. The palms BAKER , W.J. AND A.H.B. L OO . 2004. A synopsis of Micronesia and the Bonin Islands. Gentes of the genus Hydriastele (Arecaceae). Kew Herbarum 8: 423–478. Bulletin 59: 61–68. NORUP , M.V., J. D RANSFIELD , M.W. C HASE , A.S. BAKER , W.J. AND T.L.P. C OUVREUR . 2012. BARFOD , E.S. F ERNANDO AND W.J. B AKER . 2006. Biogeography and distribution patterns of Homoplasious character combinations and Southeast Asian palms. In D. G OWER , K. generic delimitation: a case study from the JOHNSON , J.E. R ICHARDSON , B. R OSEN , L. R ÜBER Indo-Pacific arecoid palms (Arecaceae: and S. W ILLIAMS (eds.). Biotic Evolution and Areceae). American Journal of Botany 93: Environmental Change in Southeast Asia. 1065–1080. Cambridge University Press, Cambridge. Pp. 164–190. TRUDGEN , M.S. AND W.J. B AKER . 2008. A revision of the Heterospathe elegans (Arecaceae) BAKER , W.J., R.A. M ATURBONGS , J. W ANGGAI AND complex in New Guinea. Kew Bulletin 63: G. H AMBALI . 2000. Siphokentia . Palms 44: 639–647. 175–181. ZONA , S. 1999. New perspectives on generic BAKER , W.J., M.J.E. C OODE , J. D RANSFIELD , S. limits and relationships in the DRANSFIELD , M.M. H ARLEY , P. H OFFMANN AND Ptychospermatinae (Palmae: Arecoideae). In R.J. J OHNS . 1998. Patterns of distribution of A. H ENDERSON and F. B ORCHSENIUS (Eds.). Malesian vascular plants. In R. H ALL AND J.D. Evolution, Variation and Classification of HOLLOWAY (Eds.). Biogeography and Geological Evolution of SE Asia. Backhuys, Palms. The New York Botanical Garden Press, Leiden. Pp 243–258. New York. Pp. 255–263.

BECCARI , O. 1924. Neue palmen Mikronesiens. ZONA , S. 2000. A personal history of Botanische Jahrbücher für Systematik, Drymophloeus in New Guinea. Palms 44: Pflanzengeschichte und Pflanzengeographie. 184–186. 59: 11–16. ZONA , S., J. F RANCISCO -ORTEGA , B. J ESTROW , W.J. BECCARI , O. AND R.G. P ICHI -SERMOLLI . 1955. BAKER AND C.E. L EWIS . 2011. Molecular Subfamiliae Arecoidearum palmae geronto- phylogenetics of the palm subtribe geae tribuum et generum conspectus. Ptychospermatinae (Arecaceae). American Webbia 11: 1–187. Journal of Botany 98: 1716–1726.

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