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Keytonorth-eastern Queensland Pandanus species (eastern coast, north of latitude ��� 19 �� ) 1. Drupes unilocolar.............................................................. Pandanus zea Drupes plurilocular (phalanges)............................................................2 2. Axillary buds of aerial branches developing into plantlets..................3 Axillary buds of aerial branches not developing into plantlets.............4 3. Pileus apex rounded to the stigmatic cluster, with narrow apical central sinuses restricted to the central stigmatic cluster; stigma on proximal face of the central cluster................................................. P. gemmifer Pileus flat or slightly domed at apex, with deep apical sinuses forming small often truncate pyramids on the whole surface of the apex; stigma on the edge of a slightly truncate pyramid..... P. grayorum 4. Syncarp with 20 ��� 45 phalanges; pileus free in the upper half..................................................................................................... P. conicus Syncarp with>60 phalanges; pileus free in the upper third or less......5 5. Prop roots massive, emerging from the stem at up to 3 m above ground; syncarp ovoid...................................................................... P. oblatus Prop roots absent or smaller emerging from the stem at or below 1 m above ground (rarely up to 2 m); syncarp spherical to ellipsoid.......6 6. Phalanges usually longer than wide; pileus sides smooth to sulcate; ascending rootlets on main stem absent or rare, leaf bases green to white; plants growing in coastal habitats......................... P. tectorius Phalanges usually as wide as, or wider than, long; pileus sides with a rib-and-groove pattern; ascending rootlets common or dense on main stem, leaf bases orange to red; plants growing in open savanna and swamps...................................................................... 7 7. Phalanges broadly cuneate to orbicular, pileus sides with dark brown cracks; sides of each carpel smooth; plants not growing in permanent water...................................................................................... P. cookii Phalanges cuneate; pileus sides without cracks; sides of carpel sides with several ridges; plants growing in swamps................................................................................................. P. solms-laubachii, Published as part of Callmander, Martin W., Gallaher, Timothy, Buerki, Sven, Zich, Frank A. & Field, Ashley R., 2021, Pandanus grayorum (Pandanaceae), a new species endemic to north-eastern Queensland (Australia), pp. 327-335 in Australian Systematic Botany 34 on page 333, DOI: 10.1071/SB20033, http://zenodo.org/record/5643372

Callmander, Martin W., Gallaher, Timothy, Buerki, Sven, Zich, Frank A., Field, Ashley R. (2021): Pandanus grayorum (Pandanaceae), a new species endemic to north-eastern Queensland (Australia). Australian Systematic Botany 34: 327-335, DOI: 10.1071/SB20033, URL: https://doi.org/10.1071/SB20033

Pandanusgrayorum Callm., Buerki & Gallaher, sp. nov. (Fig. 1.) Type: AUSTRALIA. Queensland: Cook District, Jubilee Road, off Flying Fish Point Road, Innisfail, 17 ° 290460 S, 146 ° 020260 E, 8 m, 4.x.2018, fr., B. Gray 9976 (holo: G 00446800! [3 sheets + carpological specimen]; iso: BISH!, BRI, CANB, CNS 148706! [3 sheets + carpological specimen]). Pandanus sp. (Russell River) in W.E.Cooper & W.T.Cooper, Fruits Austral. Trop. Rainforest 390 (2004). Diagnosis Pandanus grayorum is distinguished from P. gemmifer in having larger crown leaves (210 – 280 × 7 – 8 cm v. 170 – 230 × 5 – 6 cm in P. gemmifer) and syncarps (35 – 40 × 18 – 22 cm v. 23 – 25 × 10 – 18 cm); apex of phalanges flat or slightly domed at apex, divided by apical sinuses ~ 0.5 cm deep into small, often truncate pyramids as many as there are carpels (v. rounded to the stigmatic cluster, 12 – 15 mm wide with narrow apical central sinuses restricted to the central cluster); stigma on apex or on the edge of the slightly truncate apical face of each pyramid (v. on proximal face of the central apex cluster). Description Tree up to 9 m tall, stem with conical nodules, prop roots at base up to 50 cm high, upper branches (and sometimes stem) producing axillary sprouts (plantlets). Leaves 210 – 280 cm long, 7 – 8 cm wide in the middle, ~ 9 cm wide near the sheath, gradually attenuate in the upper part, coriaceous; longitudinal veins visible on both surfaces; marginal prickles beginning at 7 – 9 cm above the base and extending to the apex, antrorse, 5.5(– 6) mm in the lower third, 5(– 10) mm apart, to 2 – 5 mm long in the mid-third; 3 – 5 mm apart, up to ~ 1 mm long in the distal third, 2 – 3 mm apart; midrib armed in the upper 1/4, prickles 4 – 5 mm long), irregularly disposed (3 – 10 mm) in the middle, then regularly disposed, spaced (2 – 5 mm) as long as on the marginal in the distal 1/3; sheath 5 – 9 cm long, ~ 9 cm wide at apex, ~ 10 – 11 cm wide at base. Infrutescence terminal, a solitary syncarp on a curved peduncle; syncarp 35 – 40 × 18 – 22 cm, oblong to obovate, bright red at maturity; peduncle ~ 60 – 70 cm long, 2 cm wide at apex, trigonous, veins visible, first bract borne just above the base of syncarp, 8 bract scars along peduncle and the remaining bracts crowded at the base of the syncarp. Phalanges, 60 – 80 per syncarp, 6 – 8(– 9) cm high, 4 – 5(– 6) cm wide, 2.5 – 3.5 cm thick, 4 – 6 angled; carpels (8 –) 12 – 18(– 20), incompletely united, free in the upper ~1/3 – 1/2, tapering to the base; pileus flat or slightly domed at the apex, divided by apical sinuses ~ 0.5 cm deep into small, often truncate pyramidsasmanyastherearecarpels. Stigma 1 percarpel,oblique to vertical, generally reniform, sometimes deltoid on apex or on the edge of the slightly truncate apical face of each pyramid; endocarpbony, inlower 2/3, marginsup to 5 mm wide, 1.2 – 2.5 cm long in the centre, 1 – 2 cm wide; seed locule oblong, 1.5 – 2.5 × 0.5 – 1 cm, superior and inferior mesocarp thick and fibrous. Staminate inflorescence terminal, pendent, creamy white, up to 76 cm long, a three-sided raceme of spikes; peduncle fleshy, flaccid and soon disintegrating; lowest floral bract 67 × 4.8 cm, margins finely toothed, apex long-subulate, median bract 31 × 4.2 cm, with fine marginal prickles, basal floral bracts extending longer than distal floral bracts; spikes dense, cylindric, 6 – 10 × 2.5 – 4 cm, with manystaminate flowers; each spike of a fleshy central axis with stemonophore 10 – 13 mm longeach bearing 20 – 26 stamens; stamens with filament 1 – 2 mm long, anthers 2.5 – 4.5 × 0.8 – 1.2 mm, including an apiculus 0.3 – 0.5 mm. (Fig. 1.) Distribution This species is known from the Wet Tropics of north-eastern Queensland on and near the banks of the lower reaches of the Mulgrave, Russell, Alice, Johnstone and Moresby rivers and associated subcoastal flood plains (Fig. 2). Plants identified as P. grayorum have been observed in the Ella Bay, Moresby Range and Russell River National Parks and Etty Bay Conservation Reserve but most of the distribution occurs in fragmented remnants among agricultural land, suggesting its previous distribution may have been wider. It occurs in sympatry with P. solms-laubachii, but is parapatric with P. gemmifer, which occurs in the mid-upper reaches of the adjacent Barron, Tully and Herbert River systems to the north and south, on the Atherton Tableland, and on the McIlwraith Range on Cape York Peninsula. Habitat and ecology Pandanus grayorum occurs in lowland rainforest, specifically simple to complex mesophyll vine forest, riparian gallery forest and rainforest margins with mangrove estuaries or Melaleuca- dominated woodlands. It occurs near large river banks and associated small creeks, moist gullies and tidal estuaries near sea level. Reproduction is both via seed and from vegetative propagules, which form a localised colony of trees. The vegetative propagules float and can be dispersed in fresh and tidal waters. They are covered in sharp curved hooks and are potentially zoochorous. Pandanus grayorum can occur as a subcanopy, canopy or in open areas as an emergent tree. Phenology Flowering occurs in October to January, with fruit developing over 1 year to ripen the following October through December. Field observations suggest that there are locally male- or female-biased stands, with an overall female bias across the whole population. Vegetative propagules are produced all-year round. Conservation status Pandanus grayorum is known from few collections from several populations along rivers banks and small creeks. On the basis of collection and observation records, it has an extent of occurrence (EOO) of 231 km 2 and an area of occupancy (AOO) of 28 km 2. Most records are from localities outside national parks. No threats were observed at the known localities; however, adjacent areas are road reserves and agricultural landscapes where disturbance and clearing has occurred and may continue into the future, so it is conceivable that populations may be damaged by future activities. A preliminary assessment suggests the IUCN rating of Vulnerable [VU B1ab(iii, iv)+B2ab(iii, iv)], using the IUCN Red List Categories and Criteria (International Union for Conservation of Nature 2012). Etymology This new species is named in honour of Joy and Bruce Gray. Bruce was formerly a technical officer with CSIRO in Atherton and has collected extensively in remote areas of northern Queensland. Over his career, Bruce has made a significant contribution to the knowledge of the northern Queensland flora through his herbarium collections, discovered many new species, and made the type collections of 114 species, in addition to the one described here. Joy and Bruce welcomed the first author with a wonderful hospitality in Atherton in 2017 and Joy’ s support of Bruce has in effect enabled so many of Bruce’ s endeavours, including an epic trip to the tip of Cape York Peninsula in 2017. The first author finally dedicates this new Pandanus species for Bruce’ s 80th birthday. Notes Vegetative propagation in the genus Pandanus has been recorded in Australia and Madagascar: ‘small branchlets develop near (the) apex of branches, then drop off; these act as vegetative propagules’ (Stone 1970, p. 127). This vivipary is known in three species in Madagascar (Guillaumet 1972), but it remains rare. The active vegetative propagation in P. grayorum and P. gemmifer is so vigorous on aerial branches that they can easily be identified in the field. Fig. 2. Distribution map of Pandanus grayorum and P. gemmifer in north-eastern Queensland. Pandanus grayorum occurs in close parapatry with P. gemmifer (Fig. 2, 3), with P. grayorum occurring on the lower reaches of the Mulgrave, Russell, Alice, Johnstone and Moresby river systems, whereas P. gemmifer occurs on the Barron River system to the north and the Tully and Herbert river systems and nearby drainages to the south (Fig. 2). The two species are distinguished by the dimensions of their leaves and syncarps and the morphology of their phalanges (Table 1). Pandanus grayorum occurs in close sympatry with P. solmslaubachii (Fig. 4) and shares a somewhat similar tree architecture. Pandanusgrayorum is readily identified by the presence of axillary buds developing into plantlets, which are absent in P. solms-laubachii. Pandanus grayorum also lacks the dense ascending rootlets present in P. solms-laubachii. The two species aredistinguished by thedimensions of their leaves, shape of syncarps and the morphology of their phalanges (Table 1, Fig. 1, 4). The leaf form of P. grayorum is more arcuate and the leaves are glossy, whereas in P. solms-laubachii the leaves are more straight and folded and are moderately glaucous. The genetic and morphological similarities among P. gemmifer, P. grayorum and P. solms-laubachii suggest that hybrid speciation may have played a role in the diversification of these species (Gallaher et al. 2015). Pandanus grayorum is the second taxon to be described that shares characteristics of both Pandanus section Austrokeura and Australibrassia (however, these sections were not retrieved as monophyletic groups in the molecular phylogeny, which might suggest that these characters evolved independently in this group; Buerki et al. 2012; Gallaher et al. 2015). Both phylogenomic and population genetic studies, coupled with field observations, are recommended to clarify species delimitations within this challenging group of Pandanaceae. Additional specimen examined Australia. Queensland: Harveys Creek, swamp forest, 17 °15035.0000 S, 145 °55014.0000 E, 23.IX.1971, S.T. Blake 23631 (BRI, CANB); Bellenden Kerlanding, Russell River NP, 17 °1602200 S, 145 °5605100 E, 5 m,fr., A.Field & F. Zich 5015 (BRI, CNS, G); Polly Creek, Garadunga, in road reserve near junction of Polly Creek road and Jubilee Road, 17 °2900200 S, 146 °0200100 E, 12 m, fr., A. Field & F. Zich 5017 (BRI, CNS, G); Nind’ s Creek, Coquette Point, in road reserve of Coquette Point road abutting Moresby Range NP, 17 ° 3200200 S, 146 °0304200 E, 1 – 2 m, fr., A. Field & F. Zich 5018 (BRI, CNS, G); Moresby River tributary, in road reserve of Bruce Highway between Danesi Road and Croatto Road, 17 °3605400 S, 146 °0105000 E, 9 m, staminate fl., A. Field & F. Zich 5019 (BRI, CNS, G); Etty Bay, 17 ° 330 S, 146 °050 E, 7.XII.2011, fr., T. Gallaher 337 (BISH, HAW); Russell River, below Russell River boat camp, 17 °1602100 S, 145 ° 5605400 E, 7 m, 13. XII.2017, fr., B. Gray 9955 (BRI, CNS, G); Russell River NP, Russell River road, 17 ° 1602800 S, 145 °5605000 E, 3 m, 29.X.2003, fr., B. Gray 8831 (BRI, CANB, CNS).

Pandanusgrayorum Callm., Buerki & Gallaher, sp. nov. (Fig. 1.) Type: AUSTRALIA. Queensland: Cook District, Jubilee Road, off Flying Fish Point Road, Innisfail, 17 �� 290460 S, 146 �� 020260 E, 8 m, 4.x.2018, fr., B. Gray 9976 (holo: G 00446800! [3 sheets + carpological specimen]; iso: BISH!, BRI, CANB, CNS 148706! [3 sheets + carpological specimen]). Pandanus sp. (Russell River) in W.E.Cooper & W.T.Cooper, Fruits Austral. Trop. Rainforest 390 (2004). Diagnosis Pandanus grayorum is distinguished from P. gemmifer in having larger crown leaves (210 ��� 280 �� 7 ��� 8 cm v. 170 ��� 230 �� 5 ��� 6 cm in P. gemmifer) and syncarps (35 ��� 40 �� 18 ��� 22 cm v. 23 ��� 25 �� 10 ��� 18 cm); apex of phalanges flat or slightly domed at apex, divided by apical sinuses ~ 0.5 cm deep into small, often truncate pyramids as many as there are carpels (v. rounded to the stigmatic cluster, 12 ��� 15 mm wide with narrow apical central sinuses restricted to the central cluster); stigma on apex or on the edge of the slightly truncate apical face of each pyramid (v. on proximal face of the central apex cluster). Description Tree up to 9 m tall, stem with conical nodules, prop roots at base up to 50 cm high, upper branches (and sometimes stem) producing axillary sprouts (plantlets). Leaves 210 ��� 280 cm long, 7 ��� 8 cm wide in the middle, ~ 9 cm wide near the sheath, gradually attenuate in the upper part, coriaceous; longitudinal veins visible on both surfaces; marginal prickles beginning at 7 ��� 9 cm above the base and extending to the apex, antrorse, 5.5(��� 6) mm in the lower third, 5(��� 10) mm apart, to 2 ��� 5 mm long in the mid-third; 3 ��� 5 mm apart, up to ~ 1 mm long in the distal third, 2 ��� 3 mm apart; midrib armed in the upper 1/4, prickles 4 ��� 5 mm long), irregularly disposed (3 ��� 10 mm) in the middle, then regularly disposed, spaced (2 ��� 5 mm) as long as on the marginal in the distal 1/3; sheath 5 ��� 9 cm long, ~ 9 cm wide at apex, ~ 10 ��� 11 cm wide at base. Infrutescence terminal, a solitary syncarp on a curved peduncle; syncarp 35 ��� 40 �� 18 ��� 22 cm, oblong to obovate, bright red at maturity; peduncle ~ 60 ��� 70 cm long, 2 cm wide at apex, trigonous, veins visible, first bract borne just above the base of syncarp, 8 bract scars along peduncle and the remaining bracts crowded at the base of the syncarp. Phalanges, 60 ��� 80 per syncarp, 6 ��� 8(��� 9) cm high, 4 ��� 5(��� 6) cm wide, 2.5 ��� 3.5 cm thick, 4 ��� 6 angled; carpels (8 ���) 12 ��� 18(��� 20), incompletely united, free in the upper ~1/3 ��� 1/2, tapering to the base; pileus flat or slightly domed at the apex, divided by apical sinuses ~ 0.5 cm deep into small, often truncate pyramidsasmanyastherearecarpels. Stigma 1 percarpel,oblique to vertical, generally reniform, sometimes deltoid on apex or on the edge of the slightly truncate apical face of each pyramid; endocarpbony, inlower 2/3, marginsup to 5 mm wide, 1.2 ��� 2.5 cm long in the centre, 1 ��� 2 cm wide; seed locule oblong, 1.5 ��� 2.5 �� 0.5 ��� 1 cm, superior and inferior mesocarp thick and fibrous. Staminate inflorescence terminal, pendent, creamy white, up to 76 cm long, a three-sided raceme of spikes; peduncle fleshy, flaccid and soon disintegrating; lowest floral bract 67 �� 4.8 cm, margins finely toothed, apex long-subulate, median bract 31 �� 4.2 cm, with fine marginal prickles, basal floral bracts extending longer than distal floral bracts; spikes dense, cylindric, 6 ��� 10 �� 2.5 ��� 4 cm, with manystaminate flowers; each spike of a fleshy central axis with stemonophore 10 ��� 13 mm longeach bearing 20 ��� 26 stamens; stamens with filament 1 ��� 2 mm long, anthers 2.5 ��� 4.5 �� 0.8 ��� 1.2 mm, including an apiculus 0.3 ��� 0.5 mm. (Fig. 1.) Distribution This species is known from the Wet Tropics of north-eastern Queensland on and near the banks of the lower reaches of the Mulgrave, Russell, Alice, Johnstone and Moresby rivers and associated subcoastal flood plains (Fig. 2). Plants identified as P. grayorum have been observed in the Ella Bay, Moresby Range and Russell River National Parks and Etty Bay Conservation Reserve but most of the distribution occurs in fragmented remnants among agricultural land, suggesting its previous distribution may have been wider. It occurs in sympatry with P. solms-laubachii, but is parapatric with P. gemmifer, which occurs in the mid-upper reaches of the adjacent Barron, Tully and Herbert River systems to the north and south, on the Atherton Tableland, and on the McIlwraith Range on Cape York Peninsula. Habitat and ecology Pandanus grayorum occurs in lowland rainforest, specifically simple to complex mesophyll vine forest, riparian gallery forest and rainforest margins with mangrove estuaries or Melaleuca- dominated woodlands. It occurs near large river banks and associated small creeks, moist gullies and tidal estuaries near sea level. Reproduction is both via seed and from vegetative propagules, which form a localised colony of trees. The vegetative propagules float and can be dispersed in fresh and tidal waters. They are covered in sharp curved hooks and are potentially zoochorous. Pandanus grayorum can occur as a subcanopy, canopy or in open areas as an emergent tree. Phenology Flowering occurs in October to January, with fruit developing over 1 year to ripen the following October through December. Field observations suggest that there are locally male- or female-biased stands, with an overall female bias across the whole population. Vegetative propagules are produced all-year round. Conservation status Pandanus grayorum is known from few collections from several populations along rivers banks and small creeks. On the basis of collection and observation records, it has an extent of occurrence (EOO) of 231 km 2 and an area of occupancy (AOO) of 28 km 2. Most records are from localities outside national parks. No threats were observed at the known localities; however, adjacent areas are road reserves and agricultural landscapes where disturbance and clearing has occurred and may continue into the future, so it is conceivable that populations may be damaged by future activities. A preliminary assessment suggests the IUCN rating of Vulnerable [VU B1ab(iii, iv)+B2ab(iii, iv)], using the IUCN Red List Categories and Criteria (International Union for Conservation of Nature 2012). Etymology This new species is named in honour of Joy and Bruce Gray. Bruce was formerly a technical officer with CSIRO in Atherton and has collected extensively in remote areas of northern Queensland. Over his career, Bruce has made a significant contribution to the knowledge of the northern Queensland flora through his herbarium collections, discovered many new species, and made the type collections of 114 species, in addition to the one described here. Joy and Bruce welcomed the first author with a wonderful hospitality in Atherton in 2017 and Joy��� s support of Bruce has in effect enabled so many of Bruce��� s endeavours, including an epic trip to the tip of Cape York Peninsula in 2017. The first author finally dedicates this new Pandanus species for Bruce��� s 80th birthday. Notes Vegetative propagation in the genus Pandanus has been recorded in Australia and Madagascar: ���small branchlets develop near (the) apex of branches, then drop off; these act as vegetative propagules��� (Stone 1970, p. 127). This vivipary is known in three species in Madagascar (Guillaumet 1972), but it remains rare. The active vegetative propagation in P. grayorum and P. gemmifer is so vigorous on aerial branches that they can easily be identified in the field. Fig. 2. Distribution map of Pandanus grayorum and P. gemmifer in north-eastern Queensland. Pandanus grayorum occurs in close parapatry with P. gemmifer (Fig. 2, 3), with P. grayorum occurring on the lower reaches of the Mulgrave, Russell, Alice, Johnstone and Moresby river systems, whereas P. gemmifer occurs on the Barron River system to the north and the Tully and Herbert river systems and nearby drainages to the south (Fig. 2). The two species are distinguished by the dimensions of their leaves and syncarps and the morphology of their phalanges (Table 1). Pandanus grayorum occurs in close sympatry with P. solmslaubachii (Fig. 4) and shares a somewhat similar tree architecture. Pandanusgrayorum is readily identified by the presence of axillary buds developing into plantlets, which are absent in P. solms-laubachii. Pandanus grayorum also lacks the dense ascending rootlets present in P. solms-laubachii. The two species aredistinguished by thedimensions of their leaves, shape of syncarps and the morphology of their phalanges (Table 1, Fig. 1, 4). The leaf form of P. grayorum is more arcuate and the leaves are glossy, whereas in P. solms-laubachii the leaves are more straight and folded and are moderately glaucous. The genetic and morphological similarities among P. gemmifer, P. grayorum and P. solms-laubachii suggest that hybrid speciation may have played a role in the diversification of these species (Gallaher et al. 2015). Pandanus grayorum is the second taxon to be described that shares characteristics of both Pandanus section Austrokeura and Australibrassia (however, these sections were not retrieved as monophyletic groups in the molecular phylogeny, which might suggest that these characters evolved independently in this group; Buerki et al. 2012; Gallaher et al. 2015). Both phylogenomic and population genetic studies, coupled with field observations, are recommended to clarify species delimitations within this challenging group of Pandanaceae. Additional specimen examined Australia. Queensland: Harveys Creek, swamp forest, 17 ��15035.0000 S, 145 ��55014.0000 E, 23.IX.1971, S.T. Blake 23631 (BRI, CANB); Bellenden Kerlanding, Russell River NP, 17 ��1602200 S, 145 ��5605100 E, 5 m,fr., A.Field & F. Zich 5015 (BRI, CNS, G); Polly Creek, Garadunga, in road reserve near junction of Polly Creek road and Jubilee Road, 17 ��2900200 S, 146 ��0200100 E, 12 m, fr., A. Field & F. Zich 5017 (BRI, CNS, G); Nind��� s Creek, Coquette Point, in road reserve of Coquette Point road abutting Moresby Range NP, 17 �� 3200200 S, 146 ��0304200 E, 1 ��� 2 m, fr., A. Field & F. Zich 5018 (BRI, CNS, G); Moresby River tributary, in road reserve of Bruce Highway between Danesi Road and Croatto Road, 17 ��3605400 S, 146 ��0105000 E, 9 m, staminate fl., A. Field & F. Zich 5019 (BRI, CNS, G); Etty Bay, 17 �� 330 S, 146 ��050 E, 7.XII.2011, fr., T. Gallaher 337 (BISH, HAW); Russell River, below Russell River boat camp, 17 ��1602100 S, 145 �� 5605400 E, 7 m, 13. XII.2017, fr., B. Gray 9955 (BRI, CNS, G); Russell River NP, Russell River road, 17 �� 1602800 S, 145 ��5605000 E, 3 m, 29.X.2003, fr., B. Gray 8831 (BRI, CANB, CNS)., Published as part of Callmander, Martin W., Gallaher, Timothy, Buerki, Sven, Zich, Frank A. & Field, Ashley R., 2021, Pandanus grayorum (Pandanaceae), a new species endemic to north-eastern Queensland (Australia), pp. 327-335 in Australian Systematic Botany 34 on pages 328-333, DOI: 10.1071/SB20033, http://zenodo.org/record/5643372, {"references":["International Union for Conservation of Nature (2012) ' IUCN Red List Categories and Criteria: version 3.1 ', 2 nd edn. (IUCN Species Survival Commission: Gland, Switzerland; and Cambridge, UK)","Stone BC (1970) Observations on the genus Pandanus in Madagascar. Botanical Journal of the Linnean Society 63, 97 - 131.","Guillaumet J-L (1972) Un procede de multiplication vegetative chez les Pandanus malgaches. Adansonia 12, 429 - 431.","Gallaher T, Callmander MW, Buerki S, Keeley SC (2015) A long distance dispersal hypothesis for the Pandanaceae and the origins of the Pandanus tectorius complex. Molecular Phylogenetics and Evolution 83, 20 - 32. doi: 10.1016 / j. ympev. 2014.11.002","Buerki S, Callmander MW, Devey DS, Chappell L, Gallaher T, Munzinger J, Haevermans T, Forest F (2012) Straightening out the screw-pines: a first step in understanding phylogenetic relationships within Pandanaceae. Taxon 61, 1010 - 1020. doi: 10.1002 / tax. 615008"]}

Pandanus kalobinonensis Callm., Razakamal. & Luino, sp. nov. (Fig. 2, 3). Holotypus: MADAGASCAR. Reg. DIANA [Prov. Antsiranana]: Fkt. Anketrabe-Belinta, massif du Kalobinono, 13��38'42"S 48��40'29"E, 740 m, 16.X.2015, fr., Luino & Razakamalala 106 (G [G00341880]!; iso-: TAN!). Haec species ab omnibus congeneris syncarpio parvo globoso solitario brevipedunculato atque drupis monocarpellatis in stigma spiniforme adaxialiter sulcatum desinentibus differt. Treelet to 3���4 m tall; stem prickly, up to 5 cm dbh. Leaves 125���160 cm long, 2���3 cm wide in the middle, 3 cm wide near the sheath, gradually attenuate in the upper third, and tapering to a flagellate apex in the distal 5 cm, coriaceous; longitudinal and transverse veins visible on both surfaces; prickles white in vivo; marginal prickles beginning at (4���) 5���6 cm above the base and extending to the apex, antrorse, 2���4 mm and placed 2���8 mm apart in the lower third, to 2 mm and 5���10 mm apart in the middle third; to Infrutescence terminal, a solitary syncarp subtended by naviculate bracts, borne on a straight short peduncle; syncarp 50 �� 60���70 mm, globose to ovo��d; peduncle straight, short, 65 mm long, c. 9 mm in diam at apex, trigonous, 3��� 4 remaining bracts, c. 5��� 7.5 cm long, positioned near the base of the syncarp, boat-shaped at base and becoming flagellate in the distal part. Drupes c. 65���140 per syncarp, (15���)20(���25) mm high, 5���7 mm wide, (3���) 4���6 mm thick, 4���6 angled; carpel 1, free in the upper fourth, tapering to the base; pileus pyramidal, somewhat rounded at apex. Stigma 1, erect or curved, spiniform, on the centre of the apical face of the pileus, brown in vivo, 3���5 mm in length, stigmatic groove abaxial, covering c. 2/3 of the length of the stigma. Endocarp 10���13 mm long in the centre, Staminate plant unknown. Etymology. ��� The species epithet refers to the Kalobinono massif and its eroded landscape of Isalo Sandstone dominated by a pair of distinctly shaped domes (fig. 1). The literal translation from the Sakalava dialect means ���princess with large breasts���. Similarly, the name Galoko refers to a local king. The two summits are sacred for the local Sakalava. According to the legend, the queen embodied in the Kalobinono massif gave birth to the nearby Manongarivo massif, fathered by the king at Galoko (SOLO et al., 2008). Distribution and ecology. ��� Pandanus kalobinonensis is currently only known from the lowland moist evergreen forest in the Kalobinono massif (northwestern Madagascar) between 420 to 740 m elevation. Conservation status. ��� Pandanus kalobinonensis is known from 2 locations from the Galoko-Kalobinono Protected Area collected at a distance of c. 1 km in lowland moist evergreen forests. The holotype collection is located well within the intact forest at 740 m, but the paratype is located at 425 m and was made only a hundred meters from recently cleared forest. This latter location could be projected to disappear in a relatively near future. Lowland moist evergreen forests were highly threatened in the massifs where Tsimihety immigrants have established settlements for several decades and are practicing slash and burn agriculture (SOLO et al., 2008) (Fig. 1). The loss of moist evergreen forest area between 2006 to 2016 has reached 1408 ha, which represents 15% of the total cover of the PA (RABENANDRASANA et al., 2018). Since mid 2018, slash and burn agriculture is restricted within the strict conservation zone, but the risk of uncontrolled pasture fire is still a threat (P. Ranirison, pers. comm.). With an area of occupancy (AOO) of 4 km ��, and despite that both collections ocur within limits of the protected area, P. kalobinonensis is therefore assigned a preliminary conservation status of ���Endangered��� [EN B1ab(ii,iii,iv,v)+2ab(ii,iii,iv,v)] using IUCN Red List Categories and Criteria (IUCN, 2012). Notes. ��� Pandanus kalobinonensis is morphologically distinct among Malagasy species of Pandanus. Its small globose solitary syncarp bearing monocarpellate drupes topped by a spiniform stigma (Fig. 2, 3) resembles several Southeast Asian Pandanus species. Several species in Madagascar were hypothesized to have morphological Indo-Malaysian affinities (STONE 1974; CALLMANDER & LAIVAO, 2003). Pandanus platyphyllus Martelli was for example placed by STONE (1974) in the Malaysian Pandanus subgenus Rykia (Vrise) B.C. Stone before staminate plants were studied by HUYNH (1977) and confirmed to belong to the Afro-Malagasy subgenus Vinsonia (Warb.) B.C. Stone. Phylogenetic analyses further demonstrated that those latter morphological affinities were most likely the result of convergent evolution since all Malagasy species of Pandanus subgenus Vinsonia formed a monophyletic clade (BUERKI et al., 2012). In this context, Pandanus kalobinonensis is therefore classified in Pandanus subgenus Vinsonia. The sectional placement of this species remains to be established, like other recently discovered morphologically isolated species in Madagascar (see CALLMANDER et al., 2008, 2009b, 2013). Paratypus. ��� MADAGASCAR. Reg. DIANA [Prov. Antsiranana]: Beramanja, Anketrabe Belinta, massif du Kalobinono, 13��38'16"S 48��40'21"E, 425 m, 29.IX.2013, fr., Razakamalala et al. 7586 (TAN)., Published as part of Callmander, Martin W., Razakamalala, Richard, Luino, Iacopo, Andriamarisoa, Roger Lala & Buerki, Sven, 2020, Novelties from the Northern Mountains Complex of Madagascar V: A new threatened Pandanus (Pandanaceae) from the Kalobinono massif, pp. 99-105 in Candollea 75 (1) on pages 100-103, DOI: 10.15553/c2020v751a10, http://zenodo.org/record/5724834, {"references":["IUCN (2012). IUCN Red List Categories and Criteria: Version 3.1. Ed. 2. IUCN Species Survival Commission, IUCN, Gland & Cambridge.","HUYNH, K. - L. (1977). L'appareil male de quelques Pandanus du sousgenre Vinsonia (Pandanaceae) et sa signification taxonomique, phylogenique et evolutive. Beitr. Biol. Pflanzen. 53: 447 - 471.","BUERKI, S., M. W. CALLMANDER, D. S. DEVEY, L. CHAPPELL, T. GALLAHER, J. MUNZINGER, T. HAEVERMANS & F. FOREST (2012). Straightening out the screw-pines: a first step in understanding phylogenetic relationships within Pandanaceae. Taxon 61: 1010 - 1020.","CALLMANDER, M. W., S. BUERKI & S. WOHLHAUSER (2008). A new threatened species of Pandanaceae from northwestern Madagascar, Pandanus sermolliana. Novon 18: 421 - 424. DOI: https: // doi. org / 10.3417 / 2007101","CALLMANDER, M. W., L. GAUTIER & S. M. TRIGUI (2009 b). Pandanus nusbaumeri Callm. and L. Gaut. (Pandanaceae), a new endemic species from the Montagne d'Ambre (northern Madagascar). Candollea 64: 213 - 218.","CALLMANDER, M. W., R. BOLLIGER, R. M. HANITRARIVO & L. NUSBAUMER (2013). Pandanus tsingycola Callm. and Nusb. (Pandanaceae), a new species endemic to western Madagascar. Candollea 68: 229 - 235. DOI: https: // doi. org / 10.15553 / c 2012 v 682 a 6"]}

Pandanus papateaensis Butaud, F. Jacq & Callm., spec. nova (Fig. 3). Holotypus: POLYN��SIE-FRAN��AISE. Archipel des Tuamotu [Makatea]: Teniania (sur le plateau cot�� Est), 15��50'46"S 148��14'01"W, 70 m, 24.IV.2007, fr., Jacq, Lagouy & Huguet 1570 (PAP [PAP015953]!; iso-: G!, P!). Pandanus papateaensis Butaud, F. Jacq & Callm. can be distinguished from other species of French Polynesia by its completely fused drupes with usually 2 seminal chambers (vs 3 to more than 15), by its pileus totalling more than one third of the length (vs less than one third), and by the characteristic reddish-brown to dark-purple color of the pileus. Arbre dio��que robuste atteignant plus de 5,5 m de hauteur, stipe 12��� 16 cm de diam., droit ou rampant �� la base, aux ramifications alternes perpendiculaires au stipe et comptant jusqu����� 4 niveaux de ramification en comprenant le stipe initial, int��rieur de l�����corce verd��tre, �� racines-��chasses pr��sentes depuis la base jusqu����� c. 110 cm de hauteur sur le stipe, de 110���150 cm de long et de 4,5���5 cm de diam., couvertes d���une coiffe compos��e de grandes ��cailles sur 5 �� 7 cm, ��pines du stipe de 3���4 mm de diam��tre sur 4���6 mm de haut, ��pines des racines-��chasses de 2���2,5 mm de diam��tre sur 1,5 ���2 mm de haut. Feuilles coriaces, �� faces ventrales et dorsales vert franc, base de la face dorsale portant une tache brun-orange sur les pieds femelles; feuilles du stipe non ramifi��, de 230���250 cm de longueur, 7 cm de largeur au milieu, 8���8,5 cm pr��s de la gaine, progressivement r��tr��cies dans le tiers apical, ��pines jaun��tres �� brunes, antrorses, ��pines marginales pr��sentes �� partir de 20���27 cm au-dessus de la base jusqu����� l���apex, dans le tiers inf��rieur jusqu����� 3 mm de longueur puis devenant plus petites en allant vers l���apex (0,3���0,5 mm), espac��es de 3���7 mm tout au long de la marge, ��pines costales pr��sentes de 32���38 cm au-dessus de la base jusqu����� l���apex, gaine verd��tre de 5���6 cm de long, 13 cm de largeur apicale, 18,5���20,5 cm �� la base; feuilles des ramifications ultimes de 150���160 cm de longueur, 6��� 7 cm de largeur au milieu, 6,5��� 7,5 cm pr��s de la gaine, progressivement r��tr��cies dans le tiers apical, ��pines marginales pr��sentes �� partir de 17���23,5 cm au-dessus de la base jusqu����� l���apex, ��pines costales pr��sentes de 34���40 cm audessus de la base jusqu����� l���apex, gaine blanch��tre de 5���7 cm de long, 10 ��� 11 cm de largeur apicale, 13 ��� 14 cm �� la base; feuilles de la base de l���inflorescence m��le de 100���110 cm de longueur, 6 cm de largeur au milieu, 7 cm pr��s de la gaine, progressivement r��tr��cies dans le tiers apical, ��pines marginales pr��sentes �� partir de 20���22 cm au-dessus de la base jusqu����� l���apex, ��pines costales pr��sentes �� 32 cm au-dessus de la base jusqu����� l���apex, gaine blanche indistincte. Inflorescences m��les odorantes, d��velopp��es sur la ramification du quatri��me ordre, sous la forme d���un rac��me d�����pis long de 59���78 cm hors bract��es; p��doncules charnus de 13���16 mm de diam��tre vers la base; bract��es foliac��es jaun��tres et naviculaires au nombre de 15, dot��es d�����pines marginales et caudales courtes et souples, bract��es proximales 70 ��� 125 �� 5,5��� 7 cm au tiers inf��rieur, bract��es distales (11���)18���34 �� (1,5���) 2���4,5 cm au tiers inf��rieur; ��pis au nombre de 15, subcylindriques �� subconiques distalement, (3,5���)6���10 �� (2���) 3���4 cm, d��croissant distalement, compos��s de fascicules d�����tamines, les deux ��pis basaux dot��s d���un p��doncule de 2,5���7 cm, les autres subsessiles; fascicules blanch��tres comprenant une colonne de 5 ��� 9 mm de longueur libre d�����tamines pour 11���15 mm de longueur totale, sur 1���1,5 mm de diam��tre et portant 32���38 ��tamines; filets libres longs de 3���4 mm; anth��res jaunes p��les longues de 2���2,5 mm sur 0,3 ��� 0,4 mm de large. Inflorescences femelles non vues. Infrutescence monosyncarpique, terminale; syncarpe globo��de, 14���17 �� 14���16 cm, compos�� de c. 39���49 drupes; trognon ellipso��dal, 5,5���6 �� 4,5���5 cm; p��doncule triqu��tre, 48 �� 1,1���1,4 cm; jusqu����� 6 cicatrices de bract��es sur le p��doncule dont bract��e apicale engainante sur 2 des 3 c��t��s du p��doncule, de 25 mm de large; drupes oblongues, anguleuses, totalement fusionn��es, d��pourvues de sillon, (4,7���)5,8���6,3 �� (3,2���)4,2���5,1 �� (2,2���) 2,9 ���3,7 cm, libres sur 1/3 �� 1/2 �� l���apex, partie conn��e jaune �� orange in vivo; pileus en forme de d��me, craquel�� �� rainures longitudinales, �� c��tes saillantes, 2���2,9 cm de hauteur, brun rouge��tre �� pourpre fonc�� in vivo; plateforme stigmatique de 1,2���1,9 �� 0,7���1,3 cm; (1���)2(��� 4) stigmates r��niformes �� l���apex du pileus, c. 3,5���5 �� 2,5���4 mm; drupe biloculaire comprenant g��n��ralement 2 loges s��minales non soud��es (rarement 1, 3 ou 4 sur des drupes de petites dimensions ou composites); endocarpe obovale, turbin��, de 1,7���1,9 cm de longueur axiale sur 1���1,2 cm de largeur, �� apex distant de 1,1���1,4 cm de la base des stigmates, �� base distante de 2,2���2,7 cm de la base de la drupe; loge s��minale obovo��de, 1,2���1,5 �� 0,8���0,9 cm, �� centre supra-m��dian, m��socarpe fibreux. Etymologie. ��� L�����pith��te sp��cifique est bas�� sur ��Papatea��, une des anciennes d��nominations de l���atoll de Makatea et signifiant en tahitien ��rocher blanc��. Nom vernaculaire. ��� Ce pandanus est connu sous le nom tahitien de ��fara vare���au�� �� Makatea, ce qui signifie pandanus au fruit violet. Il s���agit d���un nom r��cent dans la mesure o�� le mot ��vare���au�� d��rive du mot anglais ��violet��. Ph��nologie. ��� Les syncarpes matures ont ��t�� observ��s d���avril �� juillet �� Makatea. Les inflorescences m��les ont ��t�� observ��es de janvier �� f��vrier et en juin en culture �� Tahiti. Distribution et ��cologie. ��� Pandanus papateaensis n���est connu que de l���atoll de Makatea o�� trois stations ont ��t�� localis��es sur le plateau entre 70 et 90 m d���altitude, �� la base du Mont Aetia, au lieu-dit Teniania et en haut de falaise �� l���amont de la plage de Moumu (Fig. 4). Il pousse sur des substrats coralliens rocheux (��feo��) tr��s diss��qu��s et difficiles �� parcourir, recouverts de formations foresti��res basses. Il se d��veloppe au sein de la for��t ou du fourr�� �� Pandanus-Ficus ou en limite de for��t �� Homalium mouo H. St. John, en m��lange avec les arbres Pandanus tectorius, Guettarda speciosa L., Ficus prolixa G. Forst. var. prolixa , Celtis pacifica Planch., les arbustes Alyxia sp., Capparis cordifolia subsp. spinosa (Lam.) Fici et les foug��res Microsorum grossum (Langsd. & Fisch.) S.B. Andrews et Davallia solida (G. Forst.) Sw. var. solida. Il n���a pas ��t�� trouv�� dans la d��pression centrale de l���atoll, ce qui peut ��tre expliqu�� par sa possible extirpation du site en raison de l���extraction du phosphate coupl��e �� une potentielle faible capacit�� de recolonisation naturelle (dispersion des fruits et d��veloppement en milieu tr��s ouvert). Sa diss��mination est tr��s probablement assur��e par le Carpophage de la Soci��t��, Ducula aurorae Peale (1848), grand pigeon frugivore end��mique des ��les de la Soci��t�� et des Tuamotu et aujourd���hui restreint uniquement �� l���atoll de Makatea. Le crabe de cocotier Birgus latro L. (1767), les bernard-l���hermite Coenobita brevimanus Dana (1852) et C. perlatus H. Milne Edwards (1837) et les rats introduits Rattus rattus L. (1758) et R. exulans Peale (1848), tous abondants �� Makatea (POUPIN, 1996; GHESTEMME, 2013), assurent potentiellement une diss��mination des fruits de Pandanus papateaensis �� plus courte distance lorsqu���ils en consomment la pulpe, �� l���image de Cardisoma carnifex Herbst (1796) pour ceux de P. tectorius (LEE, 1985). N��anmoins, les rats pourraient ��galement en consommer les graines, ce qui limiterait la r��g��n��ration de l���esp��ce. Des pieds sont aujourd���hui en culture sur le littoral de Moumu depuis 2018 (J. Ioane, pers. comm.) mais ��galement �� Tahiti o�� un pied m��le produit �� partir de drupes est int��gr�� depuis 2010 �� la plantation conservatoire d���esp��ces menac��es du domaine de Atimaono g��r��e par la Direction de l���Environnement. Ce pied provient d���une des trois drupes germ��es sur les 16 r��colt��es �� Makatea le 1er juillet 2009 (station Teniania) et mises en germination en pot �� Tahiti 15 jours apr��s (premi��re germination au bout de 4,5 mois), les deux autres pieds ��tant morts en p��pini��re. Statut de conservation. ��� Pandanus papateaensis n���est connu que de l���atoll de Makatea dont la superficie atteint 28,6 km ��. Il n���est connu que sous la forme de trois sous-populations distantes les unes des autres de moins de 5 km et au sein desquelles pr��s de 80 pieds ont ��t�� d��nombr��s. Les nombres de pieds et de sous-populations sont probablement sous-estim��s en raison de la ressemblance de l���esp��ce �� l�����tat st��rile avec le tr��s commun P. tectorius et des difficult��s extr��mes de prospection sur le substrat karstique irr��gulier caract��ristique de l�����le. Ainsi, il serait ��galement pr��sent dans les formations naturelles de Tahiva au Sud de l�����le (G. Ah-Scha, pers. comm.). Si des plantes envahissantes (Adenanthera pavonina L., Lantana camara L., Leucaena leucocephala (Lam.) de Wit, Syzygium cumini (L.) Skeels, Tecoma stans (L.) Juss. ex Kunth) sont pr��sentes �� Makatea et peuvent entrer en comp��tition avec les pieds adultes ou les plantules de Pandanus papateaensis, et si les rats introduits peuvent limiter dans une certaine mesure la r��g��n��ration de l���esp��ce en consommant ses graines, la menace principale pour ce taxon consiste en un projet de reprise d���exploitation du phosphate r��siduel pour lequel des autorisations d���exploration ont d��j�� ��t�� accord��es par le Gouvernement de la Polyn��sie fran��aise (AVENIR MAKATEA, 2019). En effet, deux des trois stations connues sont situ��es dans l���enveloppe concern��e par le projet d���exploitation (Fig. 4), ce dernier impliquant l���extirpation de toute v��g��tation ainsi que du sol humique (banque de graines). L���h��t��rog��n��it�� des menaces pesant sur les diff��rentes sous-populations conduit �� d��finir deux localit��s, la premi��re de Aetia non menac��e par l���exploitation du phosphate r��siduel tandis que la seconde, regroupant les deux autres souspopulations, menac��e par cette exploitation. L���aire d���occupation de l���esp��ce est Pandanus papateaensis est donc provisoirement ��valu�� comme ��En danger�� [EN B1ab(i,ii,iii,iv,v)+B2ab(i,ii,iii,iv,v)+D] selon les Crit��res et les Cat��gories de la Liste Rouge de l���UICN (2012). Notes. ��� Pandanus papateaensis se distingue facilement de P. tectorius par ses drupes compl��tement fusionn��es �� pileus libre sur plus de deux tiers de sa longueur, �� rainures longitudinales et d���une couleur brun rouge��tre �� pourpre fonc�� (Fig. 3). La nouvelle esp��ce est isol��e morphologiquement en Polyn��sie orientale car il s���agit du seul taxon indig��ne poss��dant g��n��ralement deux loges s��minales par drupes, P. tectorius en comprenant 3 �� 12, P. papenooensis, P. temehaniensis et P. tamaruensis en ayant g��n��ralement plus d���une dizaine (STONE, 1988), et P. arapepe H. St. John des ��les Cook n���en poss��dant qu���une seule avec un seul stigmate central (SYKES, 2016). Pour favoriser la pr��servation de P. papateaensis, nous recommandons son placement sur la liste A des esp��ces prot��g��es du code de l���environnement de Polyn��sie fran��aise, ainsi que sa multiplication au sein de plantations conservatoires ex situ sur l�����le de Makatea et sur celle de Tahiti. Par ailleurs, afin de conserver son habitat qui consiste notamment des for��ts naturelles n���ayant pas ��t�� d��grad��es par l���extraction du phosphate, nous ne pouvons qu���inciter �� classer l���ensemble de ces formations en aire prot��g��e au sens du Code de l���Environnement (aire de gestion des habitats et des esp��ces) ou �� celui du Code de l���Am��nagement (zone prot��g��e du plan g��n��ral d���am��nagement �� r��activer de la Commune de Rangiroa dont fait partie Makatea). Un tel classement serait ��galement favorable �� la conservation d���autres taxons menac��s ou quasi menac��s vivant �� Makatea comme par exemple les oiseaux indig��nes ou end��miques de l�����le (le Carpophage de la Soci��t�� [VU], le Ptilope de Makatea (Ptilinopus coralensis chalcurus Gray (1859), [NT]) et la Rousserolle de Makatea (Acrocephalus atyphus eremus Wetmore (1919), [NT])), les plantes end��miques et/ou prot��g��es (Gossypium hirsutum var. taitense (Parl.) Roberty (indig��ne et prot��g��e), Homalium mouo (end��mique de Makatea, [NT]), Myrsine ovalis var. wilderi Fosberg & Sachet (end��mique de Makatea, [VU]), Planchonella tahitensis (Nadeaud) Pierre ex Dubard (indig��ne et prot��g��e), Pritchardia mitiaroana J. Dransf. & Y. Ehrhart (end��mique de Polyn��sie orientale et prot��g��e) et Tabernaemontana pandacaqui Poir. (indig��ne et prot��g��e)) ainsi que la malacofaune terrestre malheureusement d��j�� bien appauvrie (SARTORI et al., 2014). Paratypi. ��� POLYN��SIE-FRAN��AISE. Archipel des Tuamotu [Makatea]: Te Niania Sud Moumu, en arri��re de la falaise entre Teniania et Moumu, 15��50'29.5"S 148��14'24"W, 76 m, 14.IV.2009, fr., Butaud & Jacq 2231 (PAP [015952]); Aetia, flanc NE du sommet, 15��50'34"S 148��16'30"W, 77 m, 21.IV.2009, fr., Butaud & Jacq 2262 (PAP [PAP015951]). [Archipel de la Soci��t��, Tahiti]: Atimaono, Parcelle conservatoire de la DIREN, pied cultiv�� �� partir de graines pr��lev��es �� Makatea, 27.VI.2018, fl. ♂, Butaud & Lenoble 3701 (PAP [PAP015954]); ibid loco, 4.II.2019, fl. ♂, Butaud 3753 (G, P, PAP [PAP015955, PAP015956, PAP015957]); Arue, plantule produite �� partir d���un fruit collect�� �� Teniania en juin 2009, 18.XII.2010, fr., Jacq 890 (PAP [PAP015753]); ibid loco, 13.XI.2011, fr., Jacq 1540 (PAP)., Published as part of Butaud, Jean-Fran��ois, Jacq, Fr��d��ric & Callmander, Martin W., 2019, Pandanus papateaensis (Pandanaceae): une nouvelle esp��ce menac��e de l'atoll soulev�� de Makatea (Tuamotu, Polyn��sie fran��aise), pp. 137-144 in Candollea 74 (2) on pages 138-143, DOI: 10.15553/c2019v742a3, http://zenodo.org/record/5684550, {"references":["POUPIN, J. (1996). Crustacea Decapoda of French Polynesia (Astacidea, Palinuridea, Anomura, Brachyura). Atoll Res. Bull. 442: 1 - 114.","GHESTEMME, T. (2013). Makatea, a site of major importance for endemic birds. Biodiversity Conservation Lessons Learned Technical Series 16. CEPF & CI-Pacific, Apia.","LEE, M. A. B. (1985). The Dispersal of Pandanus tectorius by the Land Crab Cardisoma carnifex. Oikos 45: 169 - 173.","AVENIR MAKATEA (2019). Notre Projet. SAS Avenir Makatea, Paofai [https: // www. avenirmakatea. com / notre-projet /].","STONE, B. C. (1988). Notes on the genus Pandanus (Pandanaceae) in Tahiti. Bot. J. Linn. Soc. 97: 33 - 48.","SYKES, W. R. (2016). Flora of the Cook Islands. National Tropical Botanical Garden, Hawaii.","SARTORI, A. F., O. GARGOMINY & B. FONTAINE (2014). Radiation and decline of endodontid land snails in Makatea, French Polynesia. Zootaxa 3772: 1 - 68."]}

Pandanus obconicus H. St. John in Pacific Sci. 17: 481. 1963. Holotypus: THAILAND. Prov. Chanthaburi: Chanburi, Makham, 100 m, 13.I.1958, fr., Smitinand 4054 (BKF [SNF080175]!). = Pandanus bifidus H. St. John in Pacific Sci. 17: 475. 1 9 6 3. Holotypus: THAILAND . Prov. Nakhon Ratchasima: Pak Tong Chai, Nakawn Rachsima, 200 m, 27.XII.1923, fr., Kerr 8132 (BK [SN247279]!; iso-: BM [BM000512621]!, K [K000781303, K000781307, K000781308]!), syn. nov. Notes. ��� ST. JOHN (1963) mentioned the holotype (Smitinand 4054) to be held at SING, but no collection has been located there. The BKF collection bears in St. John���s handwriting the mention ��� Holotypus ��� and should therefore be considered as so. STONE (1972: 48) already corrected this oversight. We have chosen to give priority to the name P. obconicus over P. bifidus based on the quality of the type collection of P. obconicus, even if this collection is only deposited at BKF., Published as part of Callmander, Martin W. & Buerki, Sven, 2018, A new species and a new synonym in Pandanus subg. Rykia (Pandanaceae) in Thailand, pp. 115-120 in Candollea 73 (1) on page 117, DOI: 10.15553/c2018v731a12, http://zenodo.org/record/5684490, {"references":["ST. JOHN, H. (1963). Revision of the genus Pandanus Stickman, part 16. Species discovered in Thailand and Vietnam. Pacific Sci. 17: 466 - 492."]}

Pandanus voradolii Callm. & Buerki, spec. nova (Fig. 2). Holotypus: THAILAND. Prov. Bueng Kan: Tham Fun waterfall, Phu Wua WS, 18°15’52”N 103°54’21”E, 345 m, 28.III.2014, fr., Callmander et al. 1163 (BKF!; iso-: BM!, G [G00428130]!, K!, SRP-061977!). Pandanus voradolii Callm. & Buerki differs from its congeners by its rheophytic ecology, acaulescent shrubby habit with a crown of narrow, subcoriaceous and flagellate leaves; its subspherical to obovate small syncarp on a long, curved, slender peduncle with 200-280 drupes. Rheophytic shrub along stream on sand, acaulescent to 1 m tall. A crown of erect leaves, pale green to glaucous below. Leaves 100-120 cm long, 2.5-3 cm wide in the middle, 2.8-3.2 cm wide near the sheath, attenuate distally on a short flagellum (c. 5 cm long), subcoriaceous; prickles white in vivo; marginal prickles beginning at 8-10 cm above the base and extending to the apex, antrorse, 2-2.5 mm in the lower third, (6-)9-15(-20) mm apart, strong, to 1.5-2 mm in the mid third; (15-)20-25(-35) mm apart, to Infructescence lateral, monosyncarpic, pending at maturity, on a curved peduncle (rarely straight); syncarp 6-7(-8) × 6-7 cm, sub-spherical to obovate; peduncle with 5-7 caducous leafy bracts; proximal bract up to 80 x 3 cm, subcoriaceous, distal bract up to 30 x 2.5 cm; peduncle, slender, 50-65 cm long, 1-1.5 cm wide at apex, veins conspicuous. Drupes, c. 200-280 per syncarp, 20-25 mm high, 7-9 mm wide, 6-8 mm depth, 6 angled; pileus flattened to steeply pyramidal, pale green in vivo. Stigma unique, bony, bifurcate (rarely simple), brown, 5-6 x 2-3 mm; endocarp c. 10 x 4 mm; seed ellipsoid, 7-8 x 3 mm; proximal and distal mesocarp fibrous. Staminate plant unknown. Etymology. – This species epithet honours Dr. Voradol Chamchumroon from the Forest Herbarium in Bangkok (BKF). Voradol is specialized in Thai Rubiaceae (especially the genus Ixora L.; CHAMCHUMROON, 2004, 2006, 2014), but he has an overarching expertise of the Thai flora. His knowledge, enthusiasm, professionalism and dedication to the Thai flora lead to the discovery of this new species that the authors are honoured to name after him. Distribution and ecology. – Pandanus voradolii is currently only known from three collections in north-eastern Thailand where it grows along streams on sands in dry evergreen forests. Conservation status. – Pandanus voradolii is currently known from three locations occurring in the vast Thai protected area network, viz. Phu Phan National Park and Phu Wua Wildlife sanctuary. Despite of the rarity of this species, which is most likely due to under collecting, this new species does not seem to be threatened in the near future and we therefore assign it a provisional status of “Least Concern” [LC] using the IUCN Red List Criteria (IUCN, 2012). Notes. – In its acaulescent shrubby habit, this new species resembles P. obconicus. Pandanus voradolii can be nevertheless easily recognized by its subcoriaceous and narrow leaves ( 4 cm and coriaceous in P. obconicus), long and curved peduncle (50-65 cm vs 20-30 cm) and sub-spherical to obovate small syncarp (6-7(-8) × 6-7 cm vs oblong 8-14 × 5-8 cm) bearing 200-280 drupes (vs 400-500). Based on its morphology, we assign P. voradolii to Pandanus subg. Rykia. Although this subgenus is polyphyletic and requires to be re-circumscribed (BUERKI et al., 2012), the new species is assigned to the true Rykia (Pandanus subsect. Rykia sensu STONE, 1974) with the type of this section, Pandanus furcatus Roxb. STONE (1982) accepted 7 native species of Pandanus subsect. Rykia including 2 possible new species for Thailand. The first of these putative new species refers to the unpublished name “Pandanus aphitrios H. St. John” based on Hansen et al. 11208 (C [C10016345]). To our knowledge, this unpublished name should be regarded as a synonym of P. reticulosus H. St. John. The second potential new species refers to the unpublished name “Pandanus hansenii H. St. John” based on Hansen et al. 11230 (C [C10016346]). This name belongs to a complex of morphologically closely-related species occurring in the mountains in the north and north-western Thailand described by ST. JOHN (1962, 1963): P. acaulescens H. St. John, P. ligulatus H. St. John and P. penetrans H. St. John. Only P. acaulescens and P. penetrans have been currently tentatively accepted (CHAYAMARIT et al., 2014) in a first Pandanaceae checklist. Pandanus subg. Rykia is certainly the most species-rich subgenus in Thailand, but more field work in this region and nearby countries (e.g. Viet-Nam and Lao PDR) is required to establish a final taxonomic treatment. Paratypi. – THAILAND. Prov. Bueng Kan: trail to Chet Si waterfall, Phu Wua WS, 18°09’32”N 103°56’55”E, 200 m, 11.III.2016, fr., Poopath et al. 1428 (BKF). Prov. Sakhon Nakhon: Phu Phan, [16°58’48”N 103°57’00”E], 30.XI.1962, fr., Phloenchit 1984 (BKF, PH).

Pandanus voradolii Callm. & Buerki, spec. nova (Fig. 2). Holotypus: THAILAND. Prov. Bueng Kan: Tham Fun waterfall, Phu Wua WS, 18��15���52���N 103��54���21���E, 345 m, 28.III.2014, fr., Callmander et al. 1163 (BKF!; iso-: BM!, G [G00428130]!, K!, SRP-061977!). Pandanus voradolii Callm. & Buerki differs from its congeners by its rheophytic ecology, acaulescent shrubby habit with a crown of narrow, subcoriaceous and flagellate leaves; its subspherical to obovate small syncarp on a long, curved, slender peduncle with 200-280 drupes. Rheophytic shrub along stream on sand, acaulescent to 1 m tall. A crown of erect leaves, pale green to glaucous below. Leaves 100-120 cm long, 2.5-3 cm wide in the middle, 2.8-3.2 cm wide near the sheath, attenuate distally on a short flagellum (c. 5 cm long), subcoriaceous; prickles white in vivo; marginal prickles beginning at 8-10 cm above the base and extending to the apex, antrorse, 2-2.5 mm in the lower third, (6-)9-15(-20) mm apart, strong, to 1.5-2 mm in the mid third; (15-)20-25(-35) mm apart, to Infructescence lateral, monosyncarpic, pending at maturity, on a curved peduncle (rarely straight); syncarp 6-7(-8) �� 6-7 cm, sub-spherical to obovate; peduncle with 5-7 caducous leafy bracts; proximal bract up to 80 x 3 cm, subcoriaceous, distal bract up to 30 x 2.5 cm; peduncle, slender, 50-65 cm long, 1-1.5 cm wide at apex, veins conspicuous. Drupes, c. 200-280 per syncarp, 20-25 mm high, 7-9 mm wide, 6-8 mm depth, 6 angled; pileus flattened to steeply pyramidal, pale green in vivo. Stigma unique, bony, bifurcate (rarely simple), brown, 5-6 x 2-3 mm; endocarp c. 10 x 4 mm; seed ellipsoid, 7-8 x 3 mm; proximal and distal mesocarp fibrous. Staminate plant unknown. Etymology. ��� This species epithet honours Dr. Voradol Chamchumroon from the Forest Herbarium in Bangkok (BKF). Voradol is specialized in Thai Rubiaceae (especially the genus Ixora L.; CHAMCHUMROON, 2004, 2006, 2014), but he has an overarching expertise of the Thai flora. His knowledge, enthusiasm, professionalism and dedication to the Thai flora lead to the discovery of this new species that the authors are honoured to name after him. Distribution and ecology. ��� Pandanus voradolii is currently only known from three collections in north-eastern Thailand where it grows along streams on sands in dry evergreen forests. Conservation status. ��� Pandanus voradolii is currently known from three locations occurring in the vast Thai protected area network, viz. Phu Phan National Park and Phu Wua Wildlife sanctuary. Despite of the rarity of this species, which is most likely due to under collecting, this new species does not seem to be threatened in the near future and we therefore assign it a provisional status of ���Least Concern��� [LC] using the IUCN Red List Criteria (IUCN, 2012). Notes. ��� In its acaulescent shrubby habit, this new species resembles P. obconicus. Pandanus voradolii can be nevertheless easily recognized by its subcoriaceous and narrow leaves ( 4 cm and coriaceous in P. obconicus), long and curved peduncle (50-65 cm vs 20-30 cm) and sub-spherical to obovate small syncarp (6-7(-8) �� 6-7 cm vs oblong 8-14 �� 5-8 cm) bearing 200-280 drupes (vs 400-500). Based on its morphology, we assign P. voradolii to Pandanus subg. Rykia. Although this subgenus is polyphyletic and requires to be re-circumscribed (BUERKI et al., 2012), the new species is assigned to the true Rykia (Pandanus subsect. Rykia sensu STONE, 1974) with the type of this section, Pandanus furcatus Roxb. STONE (1982) accepted 7 native species of Pandanus subsect. Rykia including 2 possible new species for Thailand. The first of these putative new species refers to the unpublished name ���Pandanus aphitrios H. St. John��� based on Hansen et al. 11208 (C [C10016345]). To our knowledge, this unpublished name should be regarded as a synonym of P. reticulosus H. St. John. The second potential new species refers to the unpublished name ���Pandanus hansenii H. St. John��� based on Hansen et al. 11230 (C [C10016346]). This name belongs to a complex of morphologically closely-related species occurring in the mountains in the north and north-western Thailand described by ST. JOHN (1962, 1963): P. acaulescens H. St. John, P. ligulatus H. St. John and P. penetrans H. St. John. Only P. acaulescens and P. penetrans have been currently tentatively accepted (CHAYAMARIT et al., 2014) in a first Pandanaceae checklist. Pandanus subg. Rykia is certainly the most species-rich subgenus in Thailand, but more field work in this region and nearby countries (e.g. Viet-Nam and Lao PDR) is required to establish a final taxonomic treatment. Paratypi. ��� THAILAND. Prov. Bueng Kan: trail to Chet Si waterfall, Phu Wua WS, 18��09���32���N 103��56���55���E, 200 m, 11.III.2016, fr., Poopath et al. 1428 (BKF). Prov. Sakhon Nakhon: Phu Phan, [16��58���48���N 103��57���00���E], 30.XI.1962, fr., Phloenchit 1984 (BKF, PH)., Published as part of Callmander, Martin W. & Buerki, Sven, 2018, A new species and a new synonym in Pandanus subg. Rykia (Pandanaceae) in Thailand, pp. 115-120 in Candollea 73 (1) on pages 117-119, DOI: 10.15553/c2018v731a12, http://zenodo.org/record/5684490, {"references":["CHAMCHUMROON, V. (2004). Notes on two Ixora species, new records for Thailand. Thai For. Bull., Bot. 31: 7 - 12.","CHAMCHUMROON, V. (2006). A checklist of the genus Ixora L. (Rubiaceae) in Thailand. Thai For. Bull., Bot. 34: 4 - 24","CHAMCHUMROON, V. (2014). Two new species of Ixora (Rubiaceae) from Thailand. Thai For. Bull., Bot. 42: 85 - 90.","IUCN (2012). IUCN Red List Categories and Criteria: Version 3.1. Ed. 2. IUCN Species Survival Commission, Gland & Cambridge.","BUERKI, S., M. W. CALLMANDER, D. S. DEVEY, L. CHAPELL, T. GALLAHER, J. MUNZINGER, T. HAEVERMANS & F. FOREST (2012). Straightening out the screw-pines: a first step in understanding phylogenetic relationships within Pandanaceae. Taxon 61: 1010 - 1020.","STONE, B. C. (1974). Towards an improved infrageneric classification in Pandanus (Pandanaceae). Bot. Jahrb. Syst. 94: 459 - 540.","STONE, B. C. (1982). Revisio Pandanacerum part II. Pandanus subgenus Rykia (De Vr.) Kurz (sections Gressittia, Rykiopsis, and Solmsia excepted). Fed. Mus. J. 28.","CHAYAMARIT, K., R. POOMA & N. PATTHARAHIRANTRICIN (2014). A checklist of plants in Thailand 1. Office of Natural Resources and Environmental Policy and Planning, Bangkok, Thailand."]}

3. Agrostis javanica Houtt., Nat. Hist. II(13): Aanwyz. Plaat. [2], 225, tab. 90, fig. 5. 1782. Lectotypus (designated here by Veldkamp & Callmander): INDONESIA. Java: Thunberg s.n. [?] (G-PREL [G00818013]!). = Ischaemum muticum L., Sp. Pl.: 1049. 1753. Notes. ��� MERRILL (1938: 319) discussed the G-PREL specimen but did not recognise it as Houttuyn���s. A Houttuyn specimen of Agrostis javanica in L (L-908.77-91) represents Zoysia matrella (L.) Merr. (GOUDSWAARD, 1980: 170) with watermark ���IV��� (Jean Villedary). The single gathering present in the Houttuyn herbarium in G-PREL is designated here as the lectotype., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 168, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887, {"references":["MERRILL, E. D. (1938). A critical consideration of Houttuyn's new genera and new species of plants, 1773 - 1783. J. Arnold Arbor. 19: 291 - 375.","GOUDSWAARD, P. C. (1980). The genus Zoysia (Gramineae) in Malesia. Blumea 26: 169 - 175."]}

104. Stipa littorea Burm. f., Fl. Indica: 29. 1768 [cited under Stipa spinifex L. in Nat. Hist. II(13): Aanwyz. Plaat. [2], 335; also cited under Spinifex squarrosus L., in Nat. Hist. II(13): 561, tab. 92, fig. 2. 1782]. ��� Spinifex littoreus (Burm. f.) Merr. in Philipp. J. Sci. 7: 229. 1912. Lectotypus (designated here by Wijnands, Veldkamp & Callmander): INDONESIA. Java: Anon. s.n. (G-PREL [G00360104]!). Notes. ��� The only specimen of original material in the Burman herbarium is designated here as the lectotype. This collection is annotated as ���rompot laut javanis��� [���sea grass by the Javanese���] and represents a staminate plant. Houttuyn was clearly confused about this species, which he discussed twice, obviously unaware that it is dioecious. The single collection in his Ceylon herbarium [G00818158] also is a staminate plant of Spinifex littoreus ., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 190, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887

59. Massonia depressa Houtt., Nat. Hist. II (12): Aanwyz. Plaat. [3], 424, tab. 85, fig. 1. 1780. Lectotypus (designated by JESSOP, 1976: 411): (Houttuyn, Nat. Hist. II(12): tab. 85, fig. 1. 1780). Epitypus (designated here by Callmander & Veldkamp): SOUTH AFRICA: Cape, Thunberg s.n. (UPS-THUNB n�� 7991 image seen). Notes. ��� The genus Massonia Thunb. ex Houtt. was described by Houttuyn in Natuurlijke Historie (Nat. Hist. II(12): 424. 1780), with M. depressa listed as the only species. The description was based on ���A Cape plant recently from the Hortus medicus of this place [Amsterdam] thus named by Mr. Thunberg���. The illustration shows two flowers, which were given to him by its Curator Johannes Storm in 1779. Thunberg was paid by the Hortus for sending seeds and bulbs. No specimen of Massonia has been found in the Houttuyn herbarium in G-PREL. The illustration by Houttuyn must therefore be considered to be the type. Massonia Thunb. ex L. f. (LINNAEUS f., IV.1782: 27, 193) and Massonia Thunb. (THUNBERG, 10.VII.1782: 39) are later homonyms. Massonia latifolia L. f. (and Thunb.) is based on a Masson collection that has never been located (LINNAEUS f., 1782: 193; THUNBERG, 1782: 40; see JESSOP, 1976: 410-412). This species is now widely considered to be a synonym of the earlier M. depressa. JESSOP (1976: 411) only found fruiting material at UPS-THUNB and implicitly designated Houttuyn���s plate of flowers as the lectotype. A Thunberg flowering collection in UPS-THUNB, located by M. Hjertson (UPS), is designated here as the epitype., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on pages 178-179, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887, {"references":["JESSOP, J. P. (1976). Studies in the bulbous Liliaceae in South Africa 6. The taxonomy of Massonia and allied genera. J. S. African Bot. 42: 401 - 437.","THUNBERG, C. P. (1782). Nova genera plantarum 2. Uppsala."]}

6. Amaryllis dubia Houtt., Nat. Hist. II(12): Aanwyz. Plaat. [2], 181, tab. 82, fig. 1. 1780 [non A. dubia L.]. Lectotypus (designated here by Wijnands): SOUTH AFRICA: Cape, Auge s.n. [?] (G-PREL [G00818014]!). = Nerine sarniensis (L.) Herb. in Bot. Mag.: sub tab. 2124. 1820. Notes. ��� A single element of original material is present in the Cape herbarium. This collection is designated here as the lectotype., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 168, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887

37. Crinum angustifolium Houtt., Nat. Hist. II(12): Aanwyz. Plaat. [2], 165, tab. 81, fig. 3. 1780. Lectotypus (designated here by Callmander): INDONESIA. Java: Anon. s.n. (G-PREL [G00818104]!) (Fig. 3). = Polianthes tuberosa L., Sp. Pl.: 316. 1753. Notes. ��� A single element of original material is in the Java herbarium in G-PREL. This collection is a good match for the plate and is designated here as the lectotype (Fig. 3)., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 173, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887

80. Phleum indicum Houtt., Nat. Hist. II(13): Aanwyz. Plaat. [1], 198, tab. 90, fig. 2. 1782. ��� Polytrias indica (Houtt.) Veldk. in Blumea 36: 180. 1991. Lectotypus (designated here by Veldkamp): INDONESIA. Java: Anon. s.n. (G-PREL [G00096222]!). Notes. ��� The original material designated here as the lectotype is in the Java herbarium in G-PREL. See VELDKAMP (1991) for a discussion on this species., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 184, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887, {"references":["VELDKAMP, J. F. (1991). Miscellaneous notes on Southeast Asian Gramineae. VI. Blumea 36: 179 - 181."]}

95. Renealmia capensis Houtt., Nat. Hist. II(8): Aanwyz. Plaat. [2], 335, tab. 47, fig. 1. 1777. ��� Villarsia capensis (Houtt.) Merr. in J. Arnold Arbor. 19: 360. 1938. Lectotypus (designated here by Veldkamp & Callmander): SOUTH AFRICA: Cape, Thunberg s.n. (G-PREL [G00074336]!; isolecto-: G-PREL [G00818091, G00818092]!, UPS-THUNB n�� 4228 image seen). = Menyanthes capensis Thunb., Prodr. Pl. Cap.: 34. 1794. Typus: SOUTH AFRICA: Cape, Thunberg s.n. (holo-: UPS-THUNB n�� 4228 image seen; iso-: G-PREL [G00074336, G00818091, G00818092]!). Notes. ��� Original material of Renealmia capensis in Houttuyn���s Cape herbarium is designated here as the lectotype. It was very likely collected by Thunberg and also represents a duplicate of the type of the later name Menyanthes capensis Thunb. Two further specimens in the Burman herbarium in G-PREL were annotated by Thunberg as Menyanthes capensis and represent original material of both Houttuyn���s and Thunberg���s names. Menyanthes ovata L. f. (LINNAEUS f., 1782: 133) is also based on the same Thunberg collection at UPS-THUNB, but this name is superfluous, as well as the combination by VENTENAT (1803 - 1808: sub tab. 9): Villarsia ovata (L. f.) Vent. (CALLMANDER et al., 2017). The generic Renealmia Houtt. was published by Houttuyn in 1777 (Nat. Hist. II(8): 335). Houttuyn noted ���A new genus... which I... baptise with the name now available���. Up to the beginning of the 19th century botanists often regarded that a name placed in synonymy was then available for another use, a practice that is no longer allowed. The generic name Renealmia had previously been validated by LINNAEUS (1753: 286). Both are nom. rej. [against the nom. cons. of LINNAEUS f. (1782: 7)]., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 187, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887, {"references":["VENTENAT, P. - E. (1803 - 1808). Choix de plantes, dont la plupart sont cultivees dans le jardin de Cels. Paris.","CALLMANDER, M. W., O. D. DURBIN, H. W. LACK, P. BUNGENER, P. MARTIN & L. GAUTIER (2017). Etienne-Pierre Ventenat (1757 - 1808) and the gardens of Cels and Empress Josephine. Candollea 72: 87 - 132."]}

57. Lilium japonicum Thunb. ex Houtt., Nat. Hist. II(12): Aanwyz. Plaat. [2], 245, tab. 82, fig. 2. 1780. Lectotypus (designated here by Callmander & Wijnands): JAPAN: Thunberg s.n. (G-PREL [G00818143]!; isolecto-: UPS-THUNB n�� 8137 image seen). Notes. ��� Original material is in the Japan herbarium in G-PREL. The single collection there is designated here as the lectotype. A duplicate is present in UPS-THUNB., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 178, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887

9. Anthericum bulbine Houtt., Nat. Hist. II(12): Aanwyz. Plaat. [2], 353, tab. 80, fig. 3. 1780. Lectotypus (designated here by Veldkamp & Callmander): SOUTH AFRICA: Cape, Auge s.n. [?] (G-PREL [G00818006]!; probable isolecto-: G-PREL [G00818007, G00818008]!). = Pontederia stricta Burm. f., Fl. Ind. (Prodr. Fl. Cap.): 9. 1768. ��� Onixotis stricta (Burm. f.) Wijnands in Bothalia 21: 157. 1991. ��� Wurmbea stricta (Burm. f.) J.C. Manning & Vinn. in Taxon 56: 177. 2007. Lectotypus (first step designated by WIJNANDS, 1991: 157; second step designated here by Wijnands & Callmander): SOUTH AFRICA: Cape, Auge s.n. (G-PREL [G00818007]!; probable isolecto-: G-PREL [G00818006, G00818008]!). Notes. ��� The lectotype of Anthericum bulbine is in the Cape herbarium, probably collected by Auge (WIJNANDS, 1991: 157; 1992: 499). We may never be certain whether Houttuyn���s name is a later heterotypic synonym, as it is probably also based on an Auge specimen. It is not a nom. provisorium, because although he indicated that ���Possibly [!] the Cape plant will belong to one of the species mentioned���, he did accept it (MCNEILL et al., 2012: Art. 36.1). A second step lectotypification of Pontederia stricta is made here as two collections are present in the Burman herbarium in G-PREL [G00818007, G00818008]., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 169, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887, {"references":["WIJNANDS, D. O. (1991). Notes on some Monocotyledoneae described by N. L. Burman: Asparagaceae, Asphodelaceae, Colchicaceae. Bothalia 21: 157.","WIJNANDS, D. O. (1992). Burman's Prodromus Florae Capensis. Bot. J. Linn. Soc. 109: 485 - 502.","MCNEILL, J. M., F. R. BARRIE, W. R. BUCK, V. DEMOULIN, W. GREUTER, D. L. HAWKSWORTH, P. S. HERENDEEN, S. KNAPP, K. MARHOLD, J. PRADO, W. F. PRUD'HOMME VAN REINE, G. F. SMITH, J. H. WIERSEMA & N. J. TURLAND (2012). International Code of Nomenclature for algae, fungi, and plants (Melbourne Code). Regnum Veg. 154."]}

24. Bromus japonicus Thunb. ex Houtt., Nat. Hist. II(13): Aanwyz. Plaat. [2], 315, tab. 91, fig. 4. 1782. Lectotypus (designated here by Veldkamp): JAPAN: Thunberg s.n. (G-PREL [G00380028]!; isolecto-: UPS- THUNB n�� 2503 image seen). Notes. ��� The priority of the Houttuyn name has been overlooked by almost everyone, even MERRILL (1938), with the exception of a casual mention by NAKAI (1952: 136). Thunberg in MURRAY (1784: 119) described the isonym Bromus japonicus Thunb. (also cited in THUNBERG, 1784: 52, tab. 11). The type material of Houttuyn���s earlier name is in the Japan herbarium in G-PREL. P��NZES (1936: 18) cited the type of Thunberg���s isonym at UPS-THUNB. This collection is here considered as an isolectotype., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 171, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887, {"references":["MERRILL, E. D. (1938). A critical consideration of Houttuyn's new genera and new species of plants, 1773 - 1783. J. Arnold Arbor. 19: 291 - 375.","NAKAI, T. (1952). A synoptical sketch of Korean flora. Bull. Natl. Sci. Mus. Tokyo 31.","MURRAY, J. A. (1784). Systema vegetabilium ed. 14. Gottingen.","THUNBERG, C. P. (1784). Flora japonica. Leipzig.","PENZES, A. (1936). Rozsnok (Bromus) tanulmanyok. Bot. Kozlem. 33: 98 - 138."]}

100. Saccharum japonicum Houtt., Nat. Hist. II(13): Aanwyz. Plaat. [1], 146, tab. 89, fig. 1. 1782. ��� Miscanthus japonicus (Houtt.) Andersson in ��fvers. Kongl. Vetensk.-Akad. F��rh. 12: 166. 1855. Lectotypus (designated here by Veldkamp): JAPAN: Thunberg s.n. (G-PREL [G00818080]!; isolecto-: UPS- THUNB n�� 1739 image seen; LE [Herb. Trinius 24.1] image seen). Notes. ��� A specimen of original material is in the Japan herbarium in G-PREL annotated as ��� Saccharum aristatum vel biflorum���. A duplicate in the Thunberg herbarium (UPS- THUNB n�� 1739) is annotated as S. japonicum. This was originally identified by THUNBERG (1794c: 329) as S. polydactylon L. The duplicate in LE was used by TRINIUS (1820: 169) for his combination Ripidium japonicum (Houtt.) Trin. MERRILL (1938: 320) corrected the basionym author of Miscanthus japonicus from Thunberg to Houttuyn as Thunberg���s name is a later isonym., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 189, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887, {"references":["THUNBERG, C. P. (1794 c). Museum naturalium Academiae upsaliensis 16. Uppsala.","TRINIUS, C. B. VON (1820). Fundamenta agrostographiae. Vienna.","MERRILL, E. D. (1938). A critical consideration of Houttuyn's new genera and new species of plants, 1773 - 1783. J. Arnold Arbor. 19: 291 - 375."]}

78. Paspalum dissectum (L.) L., Sp. Pl. ed. 2: 81. 1762 [cited in Nat. Hist. II(13): Aanwyz. Plaat. [1], 164, tab. 89, fig. 2. 1782]. ��� Panicum dissectum L., Sp. Pl.: 57. 1753. Notes. ��� A collection probably from the Java herbarium is in L (L-904.26-22) (see VAN STEENIS & VELDKAMP, 1973: 151). The specimen on the left-hand of L-904.26-22 is the one depicted in Natuurlijke Historie. Another specimen from Java is in G-PREL [G00818066]., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on pages 182-184, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887, {"references":["VAN STEENIS, C. G. G. J. & J. F. VELDKAMP (1973). Note on Houttuyn's herbarium. Blumea 21: 151 - 152."]}

4. Aira arundinacea sensu Houtt., Nat. Hist. II(13): Aanwyz. Plaat. [2], 229, tab. 91, fig. 1. 1782 [non A. arundinacea L.]. = Panicum bisulcatum Thunb. in Nova Acta Regiae Soc. Sci. Upsal. 7: 141. 1815. Lectotypus (designated here by Veldkamp & Callmander): JAPAN: Thunberg s.n. (UPS-THUNB n�� 1817 image seen; isolecto-: G-PREL [G00818157]!, UPS-THUNB n�� 1818 image seen). Notes. ��� Houttuyn���s text here is rather complicated and one may have trouble interpreting what exactly he was trying to convey. We have concluded that in Aanwyz. Plaat. he did not propose Aira arundinacea as a name for the ���Japansch Rietgras���, but that plate 91 with an illustration of the latter had to be placed opposite (���tegenover���, Aanwyz. Plaat. [2]) or next to (���nevens���, Nat. Hist. II(13): 229. 1782) the citation of A. arundinacea (���aangehaald���, Aanwyz. Plaat. [1]). Houttuyn discussed on pp. 229-230 three species under the heading Aira arundinacea. The first is the Linnaean one from the Near East, the ���Levantsch���, now Eragrostis collina Trin. Linnaeus, cited by Houttuyn, gave the phrase name ��� AIRA panicula oblonga secunda mutica imbricata, foliis planis���, suggesting that he had seen more than just Tournefort���s phrase name ���Gramen orientale paniculatum, portulacea semine���, where these characters are not mentioned. However, there is no specimen present in the Linnaean herbarium and he could never have seen that of Tournefort. In the absence of further data, there was little that Houttuyn could have added. The second ���Rietgras���, ���Indisch���, is mentioned in a sentence that is grammatically incorrect. It would seem that it should have begun with something like ���Voor��� [in place of]. The species clearly is Aira indica L., with a somewhat complicated nomenclatural history, which is beyond the present discussion. Houttuyn said he had transferred it to the ���Trosachtig of Gekropte Panikgras���, i.e. Panicum conglomeratum L., ���Trosachtig [panikgras]��� found in Nat. Hist. II(13): 178. 1782. This is now known as Sacciolepis indica (L.) Chase. Instead, Houttuyn replaced the latter by a very beautiful ���Japansch Rietgras���, which he said had much similarity in growth with the ���Levantsch���, according to the diagnosis (���bepaling���). A description of his own Thunberg specimen in the Japan herbarium in G-PREL [G00818157] follows. No combination is proposed for it here. In the Thunberg herbarium, there are two duplicates (UPS- THUNB n�� 1817 and 1818) on which Thunberg based his Panicum bisulcatum and from which the lectotype has been selected., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 168, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887

49. Gladiolus liliaceus Houtt., Nat. Hist. II(12): Aanwyz. Plaat. [1], 55, tab. 79, fig. 2. 1780. Lectotypus (designated here by Callmander): SOUTH AFRICA: Cape, Auge s.n. [?] (G-PREL [G00818054]!). Notes. ��� A single element of original material is in the Cape herbarium in G-PREL. This collection is designated here as the lectotype., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 176, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887

38. Cyperus javanicus Houtt., Nat. Hist. II(13): Aanwyz. Plaat. [1], 67, tab. 88, fig. 1. 1782. Lectotypus (designated here by Veldkamp): INDONESIA. Java: Anon. s.n. (G-PREL [G00360105]!). Notes. ��� A single element of original material is in the Java herbarium in G-PREL. This collection is a good match for the plate and is designated here as the lectotype., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 173, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887

56. Ixia campanulata Houtt., Nat. Hist. II(12): Aanwyz. Plaat. [1], 42, tab. 78, fig. 4. 1780. Lectotypus (designated by BROWN, 1929: 1933): SOUTH AFRICA: Cape, Auge s.n. [?] (G-PREL [G00818053]!). = Ixia aristata Thunb., Diss. Ixia: 14. 1783. ��� Ixia leucantha var. aristata (Thunb.) Baker, Hand. Irid.: 162. 1892. Lectotypus (designated here by Callmander & Veldkamp): SOUTH AFRICA: Cape, Thunberg s.n. (UPS-THUNB n�� 918 image seen). Notes. ��� The type specimen of Ixia campanulata is in the Cape herbarium in G-PREL. MERRILL (1938: 327) did not recognise it as Houttuyn���s. BROWN (1929: 133) and LEWIS (1962: 126) recognised this collection as the holotype, whereas GOLDBLATT & MANNING (2016: 185) correctly accepted Brown���s mention as an implicit lectotypification. Four collections of possibly different gatherings are present in UPS-THUNB for Ixia aristata (UPS- THUNB n�� 917, n�� 918, n�� 919 and n�� 920). The better-preserved specimen is designated here as the lectotype. Several other collections of Ixia L. are in Houttuyn���s Cape herbarium: I. maculata L. (G-PREL [G008181221]), I. monadelpha D. Delaroche (G-PREL [G008181217, G008181218]), I. polystachya L. (G-PREL [G008181219]), and I. scillaris L. (G-PREL [G00818120]). Two further specimens identified by Houttuyn as I. grandiflora D. Delaroche are now recognised in Sparaxis Ker Gawl. as S. grandiflora (D. Delaroche) Ker Gawl. (G-PREL [G00818122, G00818123])., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 178, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887, {"references":["BROWN, N. E. (1929). The Iridaceae of Burman's Florae capensis prodromus. Bull. Misc. Inform. Kew 1929: 129 - 137.","MERRILL, E. D. (1938). A critical consideration of Houttuyn's new genera and new species of plants, 1773 - 1783. J. Arnold Arbor. 19: 291 - 375.","LEWIS, G. J. (1962). South African Iridaceae. The genus Ixia. J. S. African Bot. 28: 45 - 195."]}

74. Ornithogalum dubium Houtt., Nat. Hist. II(12): Aanwyz. Plaat. [2], 309, tab. 82, fig. 3. 1780. Lectotypus (designated here by Wijnands): SOUTH AFRICA: Auge s.n. [?] (G-PREL [G00818064]!). Notes. ��� Original material is present in the Cape herbarium. It is designated here as the lectotype. There is no obvious corresponding collection in UPS-THUNB., Published as part of Wijnands, Dirk Onno, Heniger, Johannes, Veldkamp, Jan Frederik, Fumeaux, Nicolas & Callmander, Martin W., 2017, The botanical legacy of Martinus Houttuyn (1720 - 1798) in Geneva, pp. 155-198 in Candollea 72 (1) on page 182, DOI: 10.15553/c2017v721a11, http://zenodo.org/record/5721887

11. Caladium nymphaeifolium Vent. in Mag. Encycl. 4: 471. 1801 [cited in Descr. Pl. Nouv.: sub tab. 30. 1801]. Lectotypus (designated here): [INDONESIA]: ��� Wali Ila ���, (Rheede, Hort. Malab. 9: tab. 22. 1689). = Colocasia esculenta (L.) Schott Notes. ��� Despite a search for original material of this species at G and P, no collection has been located. The name Caladium nymphaeifolium was validated by Ventenat in Descriptions in 1801 in the text below the description of Caladium bicolor (Ait.) Vent. with no illustration but with a clear reference to Hortus Malabricus vol. 9 (Rheede, 1689): ���une esp��ce que j���ai trouv��e dans l���herbier Jussieu, qui est figur��e dans le IX��me vol. de l���Hort. Malabar., t. 22, et �� laquelle j���ai donn�� le nom de nymphaeifolium.��� [���a species that I found in the Jussieu herbarium, which is illustrated in the ninth vol. of the Hort. Malabar., t. 22, and to which I gave the name of nymphaeifolium.���]. Until original material is located, we designate tab. 22 in Rheede (1689) as the lectotype., Published as part of Callmander, Martin W., Durbin, Olivier D., Lack, Hans-Walter, Bungener, Patrick, Martin, Pascal & Gautier, Laurent, 2017, Etienne-Pierre Ventenat (1757 - 1808) and the gardens of Cels and Empress Jos��phine, pp. 87-132 in Candollea 72 (1) on pages 98-100, DOI: 10.15553/c2017v721a8, http://zenodo.org/record/5721919, {"references":["Rheede, H. A. van (1689). Hortus Indicus Malabaricus 9. Amsterdam."]}

Benstonea fortuita Callm. & Buerki, spec. nova (Fig. 2). Typus: MALAYSIA [BORNEO]. State Sabah: Tenom Dist., 21 km from Tenom on road to Sipittang, Lumako FR, 5��00���22���N 115��51���53���E, 620 m, 9.X.2014, fr., Callmander et al. 1197 (holo-: G [G00341180]! ; iso-: BM!, SING!, SAN!) Benstonea fortuita Callm. & Buerki is distinguished among the genus by its subacaulescent habit, its leaf abruptly attenuate at the apex with a short apicule (0.5-1 cm), its ventral pleats armed, its stout acuminate style (3-4 mm), slightly curved proximally, and its solitary globose syncarp. Subacaulescent shrub Leaves glaucous, white below, 100-120 cm long, 4-5 cm wide in the middle, 3.5 cm wide near the sheath, abruptly attenuate in the upper part, shortly apiculate (c. 0.5-1 cm long), narrowing towards the base, subcoriaceous; marginal prickles beginning at 6-9 cm above the base and extending to the apex, antrorse, c. 2 mm in the lower third, 2-6 mm apart, to c. 1 mm in the mid third, 5-10 mm apart; minute in the distal third, c. 2 mm apart; ventral pleats armed in the last 25-28 cm of the leaf with minute prickles; sheath not auriculate, whitish. Infructescence terminal, monosyncarpic, erect at maturity; syncarp in the middle of the clump of leaves, pale green, subglobose, c. 6 cm in diam.; peduncle straight, 10 cm long, 0.7 cm wide at apex, trigonous, white in vivo, longitudinal veins visible, covered with 4-5 bracts scars. Drupes c. 150, 23- 25 mm high, 4-6 mm wide, c. 6 mm in depth, 5-6-angled; pileus pyramidal, pale to dark green in vivo, shrunk when dried, 8-10 mm high (excluding style). Style unique, stout acuminate, 3-4 mm long, slightly curved proximally; stigmatic groove brown in vivo, linear, positioned on abaxial side; endocarp c. 6 mm long, walls c. 0.4 mm thick; seed locule elliptic, 4 �� 3 mm; apical mesocarp fibrous, discrete, c. 4 mm long; lower mesocarp fibrous. Staminate plant unknown. Etymology. ��� This species is only known from a single population where only one fruiting individual was observed. We were very fortunate to come across this rare and discrete taxon and the species epithet reflects our feeling when we discovered this new Benstonea. Distribution and ecology. ��� Benstonea fortuita is only known by few individuals in southwestern Sabah in the lowland Dipterocarp forest at an altitude of 600 m (Fig. 1). Conservation status. ��� Despite being known from a Forest Reserve [FR], this species is considered rare and seriously threatened. The only known population is located in the southern tip of the Lumako FR in a secondary Dipterocarp forest where few dominant trees still protect the understorey due to past logging. Benstonea fortuita is assigned a preliminary status of ���Critically Endangered��� [CR B1ab(i,iii,iv,v)] following IUCN Red List Categories and Criteria (IUCN, 2012). Notes. ��� The overall morphology of the new species can be compared to: B. glaucophylla (Ridl.) Callm. & Buerki, B. parva (Ridl.) Callm. & Buerki, and B. undulifolia (Ridl.) Callm. & Buerki. All three species share wide, abruptly attenuate leaves and a solitary syncarp with B. fortuita. Nevertheless, the new species can easily be discriminated from these species by the combination of characters presented in Table 1. In their phylogeny, BUERKI et al. (2016) showed that the new species B. fortuita belongs to clade I together with the endemic Bornean B. platystigma (Martelli) Callm. & Buerki. Although both species have the same ecology (lowland humid forest), B. fortuita can easily be distinguished from B. platystigma by its habit (subacaulescent shrub, Published as part of Callmander, Martin W. & Buerki, Sven, 2016, Two new threatened species of Benstonea Callm. & Buerki (Pandanaceae) from Sabah (Borneo, Malaysia), pp. 257-263 in Candollea 71 (2) on pages 258-259, DOI: 10.15553/c2016v712a10, http://zenodo.org/record/5721713, {"references":["IUCN (2012). IUCN Red List Categories and Criteria: Version 3.1. 2 nd ed. IUCN Species Survival Commission, Gland & Cambridge.","BUERKI, S., T. GALLAHER, T. BOOTH, G. BREWER, F. FOREST, J. T. PEREIRA & M. W. CALLMANDER (2016). Biogeography and evolution of the screw-pine genus Benstonea Callm & Buerki (Pandanaceae). Candollea 71: 211 - 223. DOI: http: // dx. doi. org / 10.15553 / c 2016 v 712 a 7"]}

Benstonea serpentinica Callm. & Buerki, spec. nova (Fig. 3). Typus: MALAYSIA [BORNEO]. State Sabah: Lahad Datu Distr., Gunung Silam, 4��58���14���N 118��10���44���E, 365 m, 3.X.2014, Callmander, Buerki & Jumian 1187 (holo-: G [G00341664] !; iso-: BM!, MO!, SING!, SAN!). Benstonea serpentinica Callm. & Buerki is distinguished among the genus by its lateral infructescence, its peduncle covered with prophylls at base, its pileus peeling at maturity, its short stout style (2-3 mm), and the pale green fruit colour of its syncarp when ripe. Treelet to 5-8 m tall, 10 cm dbh, with wide cones of proproots at the base. Each clump of leaves with several lateral erect infructescences. Leaves 300-400 cm long, 5-5.5 cm wide in the middle, 6-6.5 cm wide near the sheath, gradually attenuate in the upper part, flagellate (flagellum 22- 25 cm long), coriaceous; prickles white in vivo; marginal prickles beginning at 10-12 cm above the base and extending to the apex, antrorse (rarely retrorse in the lower 1/3), 2.5-3 mm in the lower third, (3-)4-6(-7) mm apart, strong, to c. 2 mm in the mid third; (3-)4-8(-10) mm apart, to Infructescence lateral, with 7-9 syncarps pending at maturity; syncarp 5-7 x 6-8 cm, ovoid, compressed dorso-ventrally, the distal syncarps globose and smaller (not compressed); peduncle curved (rarely straight), 20-25 cm long, 1.3-1.5 cm wide at apex, slightly flattened, veins and bracts scars visible, covered with several prophylls at base; the latter 2-4 �� 14-20(-26) cm long, coriaceous, attached at the base of the peduncule. Drupes >300 per syncarp, 15-20 mm high, 4-5 mm wide, 2-4 mm in depth, 5-angled; pileus pyramidal, pale green in vivo, 3 mm high (excluding style), covered with a white waxy surface, remaining attached and peeling at maturity, revealing the connate part of the drupes. Style unique, brown in vivo, stout, straight, shortly acuminate, slightly curved at apex, 2-3 mm long; stigmatic groove linear, positioned on abaxial side; endocarp c. 5 mm long, walls c. 0.2 mm thick; seed locule elliptic, 2-3 �� 3-3.5 mm; apical mesocarp chamber large, ovoid, c. 5.5 mm long; lower mesocarp fibrous. Staminate plant unknown. Etymology. ��� The species epithet refers to the ultramafic rocks of Mt. Silam where the species was collected and is confined. Serpentine outcrops cover c. 3,500 km 2 across the Sabah state (PROCTOR et al., 1988). These patches of ultrabasic rock are home to several endemic species (see VAN DER ENT, 2014 for more details on the plant taxa and their distributions). Distribution and ecology. ��� Benstonea serpentinica is endemic from Mt. Silam on ultramafic soil in the eastern region of Sabah (Fig. 1). This mountain, which culminates at 884 m, is home to several endemic species including the recently described Ardisia silamensis Utteridge, Julius & Suzana (UTTERIDGE et al., 2014). Conservation status. ��� Populations of Benstonea serpentinica are only known from 350 to 600 m on Mt. Silam. This patch of forest on ultramafic substrate represents c. 10 km 2 and is part of the Sapagaya FR. Logged forest, palm oil plantations and roads surround nevertheless this mountain, which is under serious human pressure. The new species is therefore preliminary assessed here as ���Critically Endangered��� [CR B1ab(iii)] following IUCN Red List Categories and Criteria (IUCN, 2012). Notes. ��� Benstonea serpentinica is most similar to B. affinis, but differs by its peduncle (short, 20-25 cm, covered with several prophylls at base vs. long, 40-45 cm, with no prophylls), its smaller drupes (15-20 �� 4-5 mm vs. 30 �� 8 mm) and style (2-3 vs. 7 mm), style morphology (stout, shortly acuminate, slightly curved proximally vs. spiniform, curved proximally) and fruit colour when ripe (pale green vs. red). In their phylogeny of Benstonea, BUERKI et al. (2016) showed that the new species B. serpentinica belongs to clade IIIc together with B. affinis, which is widespread across the Sunda shelf (including the Philippines). Paratypus. ��� MALAYSIA [BORNEO]. State Sabah: Lahad Datu Distr., Gunung Silam, 4��58���01���N 118��10���28���E, 620 m, 1.X.2014, Buerki et al. 342 (BM, G, PH, SAN)., Published as part of Callmander, Martin W. & Buerki, Sven, 2016, Two new threatened species of Benstonea Callm. & Buerki (Pandanaceae) from Sabah (Borneo, Malaysia), pp. 257-263 in Candollea 71 (2) on pages 259-261, DOI: 10.15553/c2016v712a10, http://zenodo.org/record/5721713, {"references":["PROCTOR, J. Y. F. LEE, A. M. LANGLEY, W. R. C. MUNRO & T. NELSON (1988). Ecological Studies on Gunung Silam, A Small Ultrabasic Mountain in Sabah, Malaysia. I. Environment, Forest Structure and Floristics. J. Ecol. 76: 320 - 340.","VAN DER ENT, A., R. REPIN, J. SUGAU & K. M. WONG (2014). The Ultramafic Flora of Sabah. An introduction to the plant diversity on ultramafic soils. Sabah Parks & Natural History Pulications (Borneo), Kota Kinabalu.","UTTERIDGE, T. M. A., A. JULIUS & S. SABRAN (2014). Ardisia silamensis, a new ultramafic species from Borneo; studies in Malaysian Myrsinaceae II. Kew Bull. 69: 9510.","IUCN (2012). IUCN Red List Categories and Criteria: Version 3.1. 2 nd ed. IUCN Species Survival Commission, Gland & Cambridge.","BUERKI, S., T. GALLAHER, T. BOOTH, G. BREWER, F. FOREST, J. T. PEREIRA & M. W. CALLMANDER (2016). Biogeography and evolution of the screw-pine genus Benstonea Callm & Buerki (Pandanaceae). Candollea 71: 211 - 223. DOI: http: // dx. doi. org / 10.15553 / c 2016 v 712 a 7"]}

Benstonea fortuita Callm. & Buerki, spec. nova (Fig. 2). Typus: MALAYSIA [BORNEO]. State Sabah: Tenom Dist., 21 km from Tenom on road to Sipittang, Lumako FR, 5°00’22”N 115°51’53”E, 620 m, 9.X.2014, fr., Callmander et al. 1197 (holo-: G [G00341180]! ; iso-: BM!, SING!, SAN!) Benstonea fortuita Callm. & Buerki is distinguished among the genus by its subacaulescent habit, its leaf abruptly attenuate at the apex with a short apicule (0.5-1 cm), its ventral pleats armed, its stout acuminate style (3-4 mm), slightly curved proximally, and its solitary globose syncarp. Subacaulescent shrub Leaves glaucous, white below, 100-120 cm long, 4-5 cm wide in the middle, 3.5 cm wide near the sheath, abruptly attenuate in the upper part, shortly apiculate (c. 0.5-1 cm long), narrowing towards the base, subcoriaceous; marginal prickles beginning at 6-9 cm above the base and extending to the apex, antrorse, c. 2 mm in the lower third, 2-6 mm apart, to c. 1 mm in the mid third, 5-10 mm apart; minute in the distal third, c. 2 mm apart; ventral pleats armed in the last 25-28 cm of the leaf with minute prickles; sheath not auriculate, whitish. Infructescence terminal, monosyncarpic, erect at maturity; syncarp in the middle of the clump of leaves, pale green, subglobose, c. 6 cm in diam.; peduncle straight, 10 cm long, 0.7 cm wide at apex, trigonous, white in vivo, longitudinal veins visible, covered with 4-5 bracts scars. Drupes c. 150, 23- 25 mm high, 4-6 mm wide, c. 6 mm in depth, 5-6-angled; pileus pyramidal, pale to dark green in vivo, shrunk when dried, 8-10 mm high (excluding style). Style unique, stout acuminate, 3-4 mm long, slightly curved proximally; stigmatic groove brown in vivo, linear, positioned on abaxial side; endocarp c. 6 mm long, walls c. 0.4 mm thick; seed locule elliptic, 4 × 3 mm; apical mesocarp fibrous, discrete, c. 4 mm long; lower mesocarp fibrous. Staminate plant unknown. Etymology. – This species is only known from a single population where only one fruiting individual was observed. We were very fortunate to come across this rare and discrete taxon and the species epithet reflects our feeling when we discovered this new Benstonea. Distribution and ecology. – Benstonea fortuita is only known by few individuals in southwestern Sabah in the lowland Dipterocarp forest at an altitude of 600 m (Fig. 1). Conservation status. – Despite being known from a Forest Reserve [FR], this species is considered rare and seriously threatened. The only known population is located in the southern tip of the Lumako FR in a secondary Dipterocarp forest where few dominant trees still protect the understorey due to past logging. Benstonea fortuita is assigned a preliminary status of “Critically Endangered” [CR B1ab(i,iii,iv,v)] following IUCN Red List Categories and Criteria (IUCN, 2012). Notes. – The overall morphology of the new species can be compared to: B. glaucophylla (Ridl.) Callm. & Buerki, B. parva (Ridl.) Callm. & Buerki, and B. undulifolia (Ridl.) Callm. & Buerki. All three species share wide, abruptly attenuate leaves and a solitary syncarp with B. fortuita. Nevertheless, the new species can easily be discriminated from these species by the combination of characters presented in Table 1. In their phylogeny, BUERKI et al. (2016) showed that the new species B. fortuita belongs to clade I together with the endemic Bornean B. platystigma (Martelli) Callm. & Buerki. Although both species have the same ecology (lowland humid forest), B. fortuita can easily be distinguished from B. platystigma by its habit (subacaulescent shrub

Pandanus beguinii Callm. & A. P. Keim, spec. nova (Fig. 3 A-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 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 km 2, an AOO of 54 km 2, 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 B 1ab(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 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. 3 A-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. 3539 A (BO, L, MO-6444045, MO-6444047)., Published as part of Callmander, Martin W., Keim, Ary P., Buerki, Sven & Phillipson, Peter B., 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, pp. 179-195 in Candollea 70 (2) on pages 182-187, DOI: 10.15553/c2015v702a2, http://zenodo.org/record/5720986, {"references":["IUCN (2012). IUCN Red List Categories and Criteria: Version 3.1. 2 nd ed. IUCN Species Survival Commission, Gland & Cambridge.","CALLMANDER, M. W., G. E. SCHATZ, P. P. LOWRY II, M. O. LAIVAO, J. RAHARIMAMPIONONA, S. ANDRIAMBOLOLONERA, T. RAMINO- SOA & T. CONSIGLIO (2007). Application of IUCN Red List criteria and assessment of Priority Areas for Plant Conservation in Madagascar: rare and threatened Pandanaceae indicate sites in need of protection. Oryx 41: 168 - 176.","JEBB, M. (1992). A field guide to Pandanus in New Guinea, the Bismarck Archipelago & the Solomon Islands. Christensen Research Institute, Madang.","STONE, B. C. (1992). The New Guinea species of Pandanus section Maysops St. John (Pandanaceae). Blumea 37: 31 - 61","STONE, B. C. (1987). New taxa of Pandanus (Pandanaceae) from Malesia and Papuasia. Blumea 32: 427 - 441."]}

Pandanus halmaherensis Callm. & A. P. Keim, spec. nova (Fig. 2 E-F, 4 C, 8). Typus: INDONESIA. Prov. North Maluku [Halmahera Isl.]: Central, Weda Bay, along the road to Bukit Limber, 00°31’13”N 127°59’56”E, 600 m, 27.I.2013, Callmander, Haris & Mahroji 1078 (holo-: G [G00341587]! ; iso-: BO!, L!, MO-6465087!, MO-6465088!, MO-6465089!, MO-6124975 carpo!). Haec species a congeneris infructescentia pendente solitaria ingente atque drupis pileo rubro prismatico-conico non profunde bullato in stigma spinescens desinente munitis distinguitur. Tall tree to 10-20 m tall, trunk 15-25 cm in diam., with a large cone of proproots, 1-2(-5) m long, 6-7 cm in diam. Leaves linear-attenuate, mostly (180-)220-250 (-300) cm long, 5-8 cm wide, apex gradually attenuate; sub-coriaceous to coriaceous, paler green abaxially shiny axially (especially when dry); longitudinal veins visible on both surfaces; marginal prickles borne 7-11(-15) cm above base to apex, antrorse, 1.5-2.5 mm long in the lower third, (1-) 3-10 mm apart, (0.5-) 1-2 mm long in the mid-third, 2-5 mm apart, to 0.5-1 mm long in the distal third, 0.2-0.5 (-1) mm apart, subappressed to apressed in the distal third; midrib unarmed in the lower third, prickles Infructescence terminal, pendant, red to purple, the solitary syncarp ovoid, (25-) 30-35 cm long, 11-15 cm wide, peduncle (45-) 50-80 cm long, c. 3 cm thick at apex, bearing foliaceous bracts, the proximal ones c. 60 cm long, the distal one 13 cm long, crowded at peduncle apex, spreading open at maturity. Drupes very numerous, crowded, (30-) 35-40 mm long, 4-5 mm wide, 4-5 mm thick, 1-celled, 5-7-angled, linear, pileus prismatic-conic, 8-10 mm long, shiny, shallowly bullate. Stigma one, c. 6-8 mm long, spinescent, deflected toward the syncarp apex: stigmatic groove ventral (adaxial), running along the whole length of the stigma and extending in a groove on distal part of the pileus. Endocarp bony, basal, c. 8 mm long, flat at apex, lateral walls> 1 mm thick; proximal mesocarp 2-4 mm long, fibrous, distal mesocarp 15-25 mm long, sparsely fibrous, chambered, seed locule c. 5 × 3 mm, ellipsoid. Staminate inflorescence unkown. Distribution and ecology. – Pandanus halmaherensis is found at low to mid elevations (160-700 m) in primary and disturbed evergreen tropical forests. It grows on slopes and in open areas. The species is only known from the eastern and central part of Halmahera (Fig. 4C). Conservation status. – With an EOO of 178 km 2, an AOO of 45 km 2, and 4 subpopulations, none of which are situated within the protected area network, Pandanus halmaherensis is assigned a preliminary status of “Endangered” [EN B1ab(iii)+2ab(iii)] based on the IUCN Red List Categories and Criteria (IUCN, 2012). 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. However our remarks about the current inadequate knowledge of the flora of Halmahera and the possible establishment of conservation offset sites under P. beguinii above are applicable to P. halmaherensis. Notes. – Pandanus halmaherensis is notable in having a large and massive pendant infructescence and drupes with a prismatic-conic shallowly bullate pileus with a spinescent stigma, unlike any other known species of the genus. Our new species can, however, be compared to P. sarasinorum Warb., endemic to Sulawesi (see STONE [1974b] for a complete description of this species). The two species share a similar habit (tall trees with long aerial proproots) and syncarps (very numerous and crowded cylindric drupes). Pandanus halmaherensis can nevertheless be easily distinguished by the larger dimensions of its leaves (mostly 220-250 × 5-8 cm vs. c. 150 × 4 cm in P. sarasinorum), much larger drupes (mostly 35-40 × 4-5 mm vs. 12-20 × 2-3 mm) and most notably by its larger (30-35 × 11-15 cm) solitary, ovoid syncarps (vs. an inflorescence with multiple, cylindrical syncarps, each 16-17 × 3-6 cm). This new species seems to be morphologically isolated within the genus and would only fit into Pandanus sect. Rykiopsis B. C. Stone (STONE, 1974a). However, the latter section was defined based on the multiple syncarps of its type species P. sarasinorum. Paratypi. – INDONESIA. Prov. North Maluku [Halmahera Isl.]: Central Halmahera, Weda Bay, Sake West, 00°29’39”N 127°57’44”E, 218 m, 19.VI.2013, Bangun, Mahroji & Fabanyo 843 (BO, G, L, MO-6486427, MO-3762979 carpo); East Halmahera, Weda Bay, SE of Tofu Blewen camp, 00°47’55”N 128°02’06”E, 480 m, 3.II.2013, Callmander, Fabanyo & Mahroji 1114 (BO, G, K, L, MO-6451256, MO-6451257, MO-6451258, MO-6149145 carpo); East Halmahera, Weda Bay, Kao Rahai, 00°41’54”N 128°01’02”E, 693 m, 26.II.2013, Fabanyo et al. 58 (BO, G, L, MO-6199115, MO-6451255, MO-6451254 carpo); Central Halmahera, Weda Bay, 00°31’44”N 127°54’22”E, 163 m, 11.IV.2013, Gushilman, Lasut & Fabanyo 560 (BO, G, L, MO-6472415, MO-6472416, MO-6398426 carpo).

Pandanus halmaherensis Callm. & A. P. Keim, spec. nova (Fig. 2 E-F, 4 C, 8). Typus: INDONESIA. Prov. North Maluku [Halmahera Isl.]: Central, Weda Bay, along the road to Bukit Limber, 00��31���13���N 127��59���56���E, 600 m, 27.I.2013, Callmander, Haris & Mahroji 1078 (holo-: G [G00341587]! ; iso-: BO!, L!, MO-6465087!, MO-6465088!, MO-6465089!, MO-6124975 carpo!). Haec species a congeneris infructescentia pendente solitaria ingente atque drupis pileo rubro prismatico-conico non profunde bullato in stigma spinescens desinente munitis distinguitur. Tall tree to 10-20 m tall, trunk 15-25 cm in diam., with a large cone of proproots, 1-2(-5) m long, 6-7 cm in diam. Leaves linear-attenuate, mostly (180-)220-250 (-300) cm long, 5-8 cm wide, apex gradually attenuate; sub-coriaceous to coriaceous, paler green abaxially shiny axially (especially when dry); longitudinal veins visible on both surfaces; marginal prickles borne 7-11(-15) cm above base to apex, antrorse, 1.5-2.5 mm long in the lower third, (1-) 3-10 mm apart, (0.5-) 1-2 mm long in the mid-third, 2-5 mm apart, to 0.5-1 mm long in the distal third, 0.2-0.5 (-1) mm apart, subappressed to apressed in the distal third; midrib unarmed in the lower third, prickles Infructescence terminal, pendant, red to purple, the solitary syncarp ovoid, (25-) 30-35 cm long, 11-15 cm wide, peduncle (45-) 50-80 cm long, c. 3 cm thick at apex, bearing foliaceous bracts, the proximal ones c. 60 cm long, the distal one 13 cm long, crowded at peduncle apex, spreading open at maturity. Drupes very numerous, crowded, (30-) 35-40 mm long, 4-5 mm wide, 4-5 mm thick, 1-celled, 5-7-angled, linear, pileus prismatic-conic, 8-10 mm long, shiny, shallowly bullate. Stigma one, c. 6-8 mm long, spinescent, deflected toward the syncarp apex: stigmatic groove ventral (adaxial), running along the whole length of the stigma and extending in a groove on distal part of the pileus. Endocarp bony, basal, c. 8 mm long, flat at apex, lateral walls> 1 mm thick; proximal mesocarp 2-4 mm long, fibrous, distal mesocarp 15-25 mm long, sparsely fibrous, chambered, seed locule c. 5 �� 3 mm, ellipsoid. Staminate inflorescence unkown. Distribution and ecology. ��� Pandanus halmaherensis is found at low to mid elevations (160-700 m) in primary and disturbed evergreen tropical forests. It grows on slopes and in open areas. The species is only known from the eastern and central part of Halmahera (Fig. 4C). Conservation status. ��� With an EOO of 178 km 2, an AOO of 45 km 2, and 4 subpopulations, none of which are situated within the protected area network, Pandanus halmaherensis is assigned a preliminary status of ���Endangered��� [EN B1ab(iii)+2ab(iii)] based on the IUCN Red List Categories and Criteria (IUCN, 2012). 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. However our remarks about the current inadequate knowledge of the flora of Halmahera and the possible establishment of conservation offset sites under P. beguinii above are applicable to P. halmaherensis. Notes. ��� Pandanus halmaherensis is notable in having a large and massive pendant infructescence and drupes with a prismatic-conic shallowly bullate pileus with a spinescent stigma, unlike any other known species of the genus. Our new species can, however, be compared to P. sarasinorum Warb., endemic to Sulawesi (see STONE [1974b] for a complete description of this species). The two species share a similar habit (tall trees with long aerial proproots) and syncarps (very numerous and crowded cylindric drupes). Pandanus halmaherensis can nevertheless be easily distinguished by the larger dimensions of its leaves (mostly 220-250 �� 5-8 cm vs. c. 150 �� 4 cm in P. sarasinorum), much larger drupes (mostly 35-40 �� 4-5 mm vs. 12-20 �� 2-3 mm) and most notably by its larger (30-35 �� 11-15 cm) solitary, ovoid syncarps (vs. an inflorescence with multiple, cylindrical syncarps, each 16-17 �� 3-6 cm). This new species seems to be morphologically isolated within the genus and would only fit into Pandanus sect. Rykiopsis B. C. Stone (STONE, 1974a). However, the latter section was defined based on the multiple syncarps of its type species P. sarasinorum. Paratypi. ��� INDONESIA. Prov. North Maluku [Halmahera Isl.]: Central Halmahera, Weda Bay, Sake West, 00��29���39���N 127��57���44���E, 218 m, 19.VI.2013, Bangun, Mahroji & Fabanyo 843 (BO, G, L, MO-6486427, MO-3762979 carpo); East Halmahera, Weda Bay, SE of Tofu Blewen camp, 00��47���55���N 128��02���06���E, 480 m, 3.II.2013, Callmander, Fabanyo & Mahroji 1114 (BO, G, K, L, MO-6451256, MO-6451257, MO-6451258, MO-6149145 carpo); East Halmahera, Weda Bay, Kao Rahai, 00��41���54���N 128��01���02���E, 693 m, 26.II.2013, Fabanyo et al. 58 (BO, G, L, MO-6199115, MO-6451255, MO-6451254 carpo); Central Halmahera, Weda Bay, 00��31���44���N 127��54���22���E, 163 m, 11.IV.2013, Gushilman, Lasut & Fabanyo 560 (BO, G, L, MO-6472415, MO-6472416, MO-6398426 carpo)., Published as part of Callmander, Martin W., Keim, Ary P., Buerki, Sven & Phillipson, Peter B., 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, pp. 179-195 in Candollea 70 (2) on page 193, DOI: 10.15553/c2015v702a2, http://zenodo.org/record/5720986, {"references":["IUCN (2012). IUCN Red List Categories and Criteria: Version 3.1. 2 nd ed. IUCN Species Survival Commission, Gland & Cambridge.","STONE, B. C. (1974 b). Studies in Malesian Pandanaceae XII. Taxo- nomic revision of Pandanus sections Solmsia and Rykiopsis and of section Rykia subsection Gressettia, all of subgenus Rykia. Fed. Mus. J. 17: 99 - 163.","STONE, B. C. (1974 a). Towards an improved infrageneric classification in Pandanus (Pandanaceae). Bot. Jahrb. Syst. 94: 459 - 540."]}

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 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. 2 A-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 (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 (P. tectorius Parkinson, Published as part of Callmander, Martin W., Keim, Ary P., Buerki, Sven & Phillipson, Peter B., 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, pp. 179-195 in Candollea 70 (2) on pages 181-182, DOI: 10.15553/c2015v702a2, http://zenodo.org/record/5720986, {"references":["GALLAHER, T., M. W. CALLMANDER, S. BUERKI & S. C. KEELEY (2015). A long distance dispersal hypothesis for the Pandanaceae and the origins of the Pandanus tectorius complex. Molec. Phylogen. Evol. 83: 20 - 32","LIM, T. K. (2012). Edible medicinal and non-medicinal plants. Vol. 4, Fruits. Springer.","KEIM, A. P. (2009). Pandanaceae of the island of Yapen, Papua (West New Guinea), Indonesia, with their nomenclature and notes on the rediscovery of Sararanga sinuosa, and several new species and records. Blumea 54: 255 - 266.","KEIM, A. P., S. SUSIARTI & M. AMIR (2008). Taksonomi Pandanaceae Pulau Seram. In: UBAIDILLAH, R., E. B. WALUJO, A. J. ARIEF & H. JULISTIONO (ed.), Laporan Teknik Pusat Penelitian Biologi- LIPI: 1281 - 1326. LIPI, Bogor.","STONE, B. C. (1974 a). Towards an improved infrageneric classification in Pandanus (Pandanaceae). Bot. Jahrb. Syst. 94: 459 - 540.","STONE, B. C. (1992). The New Guinea species of Pandanus section Maysops St. John (Pandanaceae). Blumea 37: 31 - 61","JEBB, M. (1992). A field guide to Pandanus in New Guinea, the Bismarck Archipelago & the Solomon Islands. Christensen Research Institute, Madang.","STONE, B. C. (1982). New Guinea Pandanaceae: first approach to ecology and biogeography. In: GRESSIT, J. L. (ed.), Biogeography and ecology of New Guinea 1: 401 - 436. W. Junk Plush.","RUMPHIUS, G. E. (1741 - 1755). Herbarium amboinense. Vol. 1 - 6. Franciscus Changuion, Amsterdam."]}

Key to the Malagasy species of Xyris 1 Keel of lateral sepals entire; bracts, leaf and stem bases pale or dark brown when dried....................................... 2 1a. Keel of lateral sepals toothed, at least in the middle; bracts, leaf and stem bases always dark brown when dried....... 5 2 Scape spirally twisted, base swollen, bulbous; leaves filiform, c. 1 mm wide............................................. X. labatii 2a. Scape straight, not twisted, base not swollen; leaves linear, more than 2 mm wide.......................................... 3 3. Bracts with a distinct sub-terminal, triangular or rhomboid mark; inflorescence broadly ovoid at anthesis, or rarely subspherical; plants perennial........................................ 4 3a. Bracts without a distinct sub-terminal, triangular or rhomboid mark; inflorescence sub-spherical at anthesis; plants usually annual................................................ X. capensis 4. Keel of lateral sepals not winged; bract apices conspicuously thickened, dark brown, somewhat recurved; plant less than 15 cm tall.......................................... X. humilis 4a. Keel of lateral sepals winged; bract apices not conspicuously thickened, light brown, not recurved; plant more than 25 cm tall.......................................................... X. anceps 5. Leaves large (0.6-0.8 cm); inflorescence c. 10 mm in diam., broadly ovate-elliptic, not elongating with age................. ؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆؆. X. marojejyensis 5a. Leaves narrow ( 6. Leaves 40-75 × 0.3 cm; inflorescence ovoid-cylindrical, c. 2 cm long at anthesis.................................... X. baronii 6a. Leaves 20-40 × 0.15-0.2 cm; inflorescence ellipsoid, c. 1 cm long at anthesis................................ X. congensis

Xyris labatii Rakoton., Callm. & Phillipson, spec. nova (Fig. 3, 4). Typus: MADAGASCAR. Prov. Antananarivo: Ambatofinandrahana, massif de l���Itremo, Atsirakambiaty, 20��35���22���S 46��34���01���E, 1680 m, 27.III.1999, fl., Labat 3050 (holo-: P [P00160157]!; iso-: G [G00341191]!, K, MO!, TAN!). Haec species a congeneris madagascariensibus caule e base tumida orto, foliis filiformibus, caulibus scapisque spiraliter tortis atque seminibus obovoideis distinguitur. Clumped perennial herb, tufted (10-) 20-50 cm. Stem spirally twisted, from the bulbous base to the shoot. Leaves up to 23 �� 0.1 cm; sheaths (0.4-) 1-5 cm long, ridged, ligule c. 1 mm, slightly acute, brown-hyaline; blades filiform and coriaceous, strigillose, symmetric and aristate at the apex, margins thickened. Scapes up to (1-)8-28 �� 0.1 cm, terete. Inflorescences ellipsoid, c. 0.3-0.6 �� (3-) 0.5-0.6 cm, red-brown. Sterile bracts, 2-6, obovate, glossy brown, rounded to acute at the apex; fertile bracts ovate-elliptic c. 0.6 �� 0.3 cm, apex acute, concave, smooth glossy brown, hyaline towards the margins, sometimes erose. Flower with lateral sepals c. 1.3 �� 0.4 cm, slightly curved, keeled, hyaline and darker on the keel, smooth. Petals obovate, c. 0.1 �� 0.08 cm yellow. Stamens 3, anther sessile, linear-shaped, c. 1.2 �� 0.2 mm, epipetalous. Ovary obovate c. 3 �� 1.2 mm, brown. Style 3-branched, 1.2 mm long; stigma twisted, 2.4 mm long. Fruits obovate, 2 �� 1.5 mm, glabrous, with a longitudinal dark-brown dehiscence groove. Seeds obovoid, 0.3 �� 0.2 mm, dark brown Distribution and ecology. ��� Xyris labatii is known from the central highlands of Madagascar, in the Amoron���i Mania, Vakinankaratra and Haute Matsiatra regions. It grows on rocky areas, often on quartzite from 1200 to 2500 m (Fig. 2). Etymology. ��� The species epithet honours our friend and colleague, Jean-No��l Labat (1959-2011) who disappeared too soon. Jean-No��l collected the type specimen and as is often the case, he made a useful photograph to accompany his excellent collection. Notes. ��� Xyris labatii can be recognized by its bulbous stem bases and its twisted leaves and scapes. It is similar to X. humilis, which also occurs in the central highlands of Madagascar, but the new species can be distinguished by its filiform leaves (vs. very narrowly linear in X. humilis) (8-23 �� 0.1 cm vs. 3-5 �� 0.1-0.2 cm). Conservation status. ��� With an EOO of 12,460 km ��, and an AOO of 153 km �� and eleven subpopulations, two of which are in the protected area network (Andringitra), X labatii is assigned a preliminary status of ���Least Concern��� (LC) following IUCN Red List Categories and Criteria (IUCN, 2012) Paratypi. ��� MADAGASCAR. Prov. Antananarivo: Mont Ibity, c. 27 km SW of Antsirabe, 20��05���24���S 47��00���23���E, 1840-1890 m, 15.III.2004, Almeda 8706 (CAS, TAN); Antsirabe, Beapombo, �� 400 m avant d���arriver au sommet d���Ibity, 20��04���58���S 47��00���37���E, 2170 m, 5.II.2003, Andriamihajarivo 103 (MO, P [P00853040], TAN); ibid. loc., 20��10���S 47��03���E, 1300-1800 m, 6.III.1985, Dorr & al. 3853 (MO, P [P01641000]); ibid. loc., 20��07���00���S 47��01���00���E, 24.XII.1965, Peltier 2127 (P [P01640947]); Mahasoa �� Ambohimasina, Betafo, [19��50���S 46��51���E], III.1976, A. Rakotozafy 1736 (TAN); Hauts-plateaux, env. �� 15 km �� l���W de Arivonimamo (entre Arivonimamo et Soamahamanina, 60 km �� l���W de Antananarivo), 18��54���43���S 47��05���21���E, 1260 m, 23.I.2000, Raynal-Roques & J��r��mie 24816 (G, K, MO, P [P00853043], TAN, WAG); ibid. loc, 20��05���33���S 47��00���14���E, 2080 m, 16.II.2003, Schatz & al. 4075 (MO); ibid. loc., 20��04���10���S 47��00���16���E, 1700 m, 17.II.2003, Schatz & al. 4130 (MO, P, TAN). Prov. Fianarantsoa: Sommet tr��s rocheux (quartzitique) �� l���E d���Antsirakambiaty, �� l���W d���Ambatoandrano, 20��34���39���S 46��34���26���E, 1681 m, 18.IV.2003, Andriamihajarivo 179 (MO, P, TAN); 10 km W of Ivato on route #35, [20��34���30���S 46��37���30���E], 1500-1685 m, 27.I.1975, Croat 29532 (MO, P [P01640906], TAN); Vicinity of col de Itremo, [20��34���30���S 46��37���30���E], 1500-1685 m, 27.I.1975, Croat 29830 (MO, TAN); Env. d���Ambatofinandrahana, [20��33���S 46��48���E], 1600-1800 m, 19.II.1938, Decary 12993 (P [P00853042]); ibid. loc., Decary 13090 (P [P00853041]); Mt Boby, [22��11���S 46��53���E], 20. III.1945, Homolle 1209 (P [P01924015]); ibid. loc., Homolle 1201 (P [P01924014], TAN); Massif de l���Andringitra (Iratsy), vall��es de la Riambava et de l���Antsifotra et montagnes environnante, [22��12���S 46��55���E], 2000- 2500 m, 27. XI.1924, Humbert 3890 (P [P01640921, P01640957]); Massif de l���Itremo, W of Ambatofinandrahana: Montagnes �� l���W (W Betsileo), [20��34���30���S 46��37���30���E], 1500-1700 m, 17-22. I. & 18- 22.IV.1955, Humbert 28325 (P [P019524021]); ibid. loc., Humbert 29893 (P [P01640936]); ibid. loc., Humbert 29936 (G, K, MO, P [P00730639, P00730641], TAN, WAG); Route Ambositra �� Ambatofinandrahana sur rocher apr��s l���embranchement d���Ivato, 20��33���S 46��48���E, III.1960, Keraudren 158 (P [P01640896]); Itremo mountain, c. 1 km below the highest point of the road, 20��36���S 46��35���E, 1600 m, fl., 17. III.1995, Lye, Arnstein & Ranaivojaona 20894 (TAN); Itremo massif, W of Ambatofinandrahana, along road to Col d���Itremo, ca. 2 km before (NE of) bridge over Ambalarangolana creek, 20��33���58���S 46��35���35���E, 1450 m, 10.XI.2002, Lowry & al. 5914 (MO, TAN); ibid. loc., 20��40���S 46��35���E, 1600 m, fl., 23. III.1971, Mabberley 780 (K, TAN); ibid. loc., [20��37���S 47��12���E], 24.XII.1965, Peltier 5612 (P [P0164927]); ibid. loc., 20��36���45���S 46��35���05���E, 1650 m, 14. III.1992, Phillipson, Clement & Rafamantananantsoa 3889 (MO, TAN); ibid. loc., inselberg �� 11,7 km d���Ivato vers la route d���Ambatofinandrahana, et c. 4 km avant le village d���Anjoman��� Ankona, 20��39���45���S 47��07���42���E, 1500 m, fl., 24. III.2010, Rakotoarivelo & al. 330 (MO, P, TAN); ibid. loc., about 6 km NW of Itremo, 20��33���35���S 46��34���55���E, 1333 m, 9.II.2009, Rakotoarisoa 628 (K, MO, P, TAN)., Published as part of Rakotonirina, Nivo, Callmander, Martin W., Phillipson, Peter B. & Lock, J. Michael, 2014, Two new species of Xyris L. (Xyridaceae) endemic to Madagascar with a key to the Malagasy species, pp. 171-178 in Candollea 69 (2) on page 175, DOI: 10.15553/c2014v692a9, http://zenodo.org/record/5768228, {"references":["IUCN (2012). IUCN Red List Categories and Criteria: Version 3. 1. 2 nd edition. IUCN Species Survival Commission, Gland & Cambridge."]}

Xyris marojejyensis Lock, Rakoton., Callm. & Phillipson, spec. nova (Fig. 1). Typus: MADAGASCAR. Prov. Antsiranana: Sommet oriental du massif de Marojejy (NE) �� l���W de la haute Manantenina, affluent de la Lokoho, [14��25���48���S 49��43��� 48���E], 1850-2137 m, fl., 17-20.XII.1948, Humbert & Capuron 22770 (holo-: P [P00730642]!; iso-: BR!, G [G00406221]!, K!, MO!, P [P00730640]!, TAN!). Haec species a congeneris madagascariensibus caule robusto, foliis ca. 8 mm latis, inflorescentia nitida in vetustate non elongata atque bractis fertilibus grandibus (ca. 10 �� 8 mm) distinguitur. Large perennial herb to 1 m. Leaves up to 35 �� 0.6-0.8 cm; sheaths 5-6 cm long, weakly ridged; ligule c. 2 mm long, rounded, brown-hyaline; lamina up to 35 cm long and 0.8 cm wide, flat, thickened and coriaceous, smooth, slightly asymmetric and blunt at the apex, margins thickened. Scapes terete up to 70 cm tall and 3 mm in diam., with a single deflexedpubescent longitudinal ridge. Inflorescences broadly ellipsoid, c. 10 mm in diam. Lowest pair of sterile bracts broadly ovate to orbicular, c. 4 �� 4 mm, glossy brown, rounded at the apex; upper sterile bracts, 4-6, similar but larger; fertile bracts, 6-8, broadly ovate-elliptic, c. 10 �� 8 mm, glossy brown, slightly darker towards the rounded or retuse-mucronate apex, lacking a dorsal mark, concave, smooth. Flower lateral sepals slightly curved, strongly keeled c. 8 �� 0.6 mm, very pale brown, much darker on the keel, the keel spinose-pubescent, particularly towards the middle. Petals suborbicular, c. 1 cm in diam., yellow. Staminodia not present. Stamens 3, alternating with the petals, epipetalous; anthers sessile, linearc. 4 x 1 mm, with a longitudinal dehiscence. Ovary obovate, c. 5.5 �� 1.9 mm, brown. Style 3-branched, the branches capitate at the apex. Young fruit ovoid, 0.5 �� 0.2 mm, brown, glabrous. Seeds ovoid 0.1 �� 0.03 mm, brown, glabrous. Distribution and ecology. ��� Xyris marojejyensis is known only from between 1300 to 2100 m in the Marojejy massif in north-eastern Madagascar, growing in ericoid shrubland on substrates derived from gneiss and quartzite (Fig. 2). Notes. ��� The large size of its leaves and bracts, and the shape of its scapes and flowers easily distinguished X. marojejyensis from its Malagasy congeners. The new species can be distinguished from the other larger species occurring in Madagascar, X. baronii and X. congensis, most easily in the case of the former by the shape of its scapes (terete vs. ovoidfusiform in X. baronii) and dimensions of its bracts (c. 10 �� 8 vs. 4-5 �� 0.5 mm in X. baronii), and from the latter by the broader leaves (0.6-0.8 vs. 0.15-0.3 cm in X. congensis) and corolla (1 cm. in diam. vs. 0.35 cm in diam. in X. congensis). Conservation status. ��� With an EOO of 398 km ��, and an AOO of 63 km �� and three known subpopulations all occurring in a Protected Area (Marojejy), X. marojejyensis is assigned a preliminary status of Least Concern [LC] due to its distribution at high elevation in a non-threatened habitat. Paratypi. ��� MADAGASCAR. Prov. Antsiranana: Sommet du Marojejy, [14��25���48���S 49��43���48���E], 29.III.1947, Cours 3537 (P [P01640932], TAN); Vall��e inf��rieure de l���Androranga, affluent de la Bemarivo (NE), aux env. d���Antongondriha, massif de Betsomanga, [14��15���30���S 49��44���00���E], 1300-1950 m, 17-20.XI.1950, Humbert & Capuron 24333 (P [P01640952]); Vall��e de la Lokoho (NE), Mt Beondroka au N de Maroambihy, [14��30���00���S 49��50���30���E], 1400-1450 m, 17-22.III.1949, Humbert & Capuron 23618 (G, K, MO, P [P00853045],); ibid. loc., Humbert & Capuron 23621 (P [P0073 0621]); Along the trail to the summit of Marojejy E, NW of Mandena, 14��26���S 49��43���E, 1900-2133 m, 15.II.1989, Miller & Lowry 4153 (MO, P [P00853038], TAN); Massif de Marojejy, [14��25���40���S 49��43���48���E], 2000-2137 m, XI.1972, Morat 4081 (P [P00853037]); ibid. loc., 10.5 km NW of Manantenina, along tributary at head of Andranomifototra river, 14��26���24���S 49��44���30���E, 1625 m, 4-13.XI. 1996, Rakotomalaza & al. 869 (MO); ibid. loc., partie SW, 14��28���S- 14��37���S 49��33���E-49��42���E, 1295-1620 m, 12.V.1989, Randrianasolo 16 (MO); ibid. loc., above Manentinina Village, 14��26���S 49��43���E, 1600-2137 m, 28.III.1990, Randrianasolo 139 (MO, P [P00730622], TAN); ibid. loc., 14��25���45���S 49��42���30���E, 1672 m, 9.III.1994, Rasoavimbahoaka & al. 161 (MO, P [P00730624], TAN); ibid. loc., au sommet de Marojejy, 14��26���50���S 49��43���57���E, 2132 m, 23- 24.III.1995, Rasoavimbahoaka & al. 520 (MO, P [P00730623], TAN)., Published as part of Rakotonirina, Nivo, Callmander, Martin W., Phillipson, Peter B. & Lock, J. Michael, 2014, Two new species of Xyris L. (Xyridaceae) endemic to Madagascar with a key to the Malagasy species, pp. 171-178 in Candollea 69 (2) on page 173, DOI: 10.15553/c2014v692a9, http://zenodo.org/record/5768228

Key to the Malagasy species of Xyris 1 Keel of lateral sepals entire; bracts, leaf and stem bases pale or dark brown when dried....................................... 2 1a. Keel of lateral sepals toothed, at least in the middle; bracts, leaf and stem bases always dark brown when dried....... 5 2 Scape spirally twisted, base swollen, bulbous; leaves filiform, c. 1 mm wide............................................. X. labatii 2a. Scape straight, not twisted, base not swollen; leaves linear, more than 2 mm wide.......................................... 3 3. Bracts with a distinct sub-terminal, triangular or rhomboid mark; inflorescence broadly ovoid at anthesis, or rarely subspherical; plants perennial........................................ 4 3a. Bracts without a distinct sub-terminal, triangular or rhomboid mark; inflorescence sub-spherical at anthesis; plants usually annual................................................ X. capensis 4. Keel of lateral sepals not winged; bract apices conspicuously thickened, dark brown, somewhat recurved; plant less than 15 cm tall.......................................... X. humilis 4a. Keel of lateral sepals winged; bract apices not conspicuously thickened, light brown, not recurved; plant more than 25 cm tall.......................................................... X. anceps 5. Leaves large (0.6-0.8 cm); inflorescence c. 10 mm in diam., broadly ovate-elliptic, not elongating with age................. ��������������������������������������������������������������. X. marojejyensis 5a. Leaves narrow (X. baronii 6a. Leaves 20-40 �� 0.15-0.2 cm; inflorescence ellipsoid, c. 1 cm long at anthesis................................ X. congensis, Published as part of Rakotonirina, Nivo, Callmander, Martin W., Phillipson, Peter B. & Lock, J. Michael, 2014, Two new species of Xyris L. (Xyridaceae) endemic to Madagascar with a key to the Malagasy species, pp. 171-178 in Candollea 69 (2) on pages 172-173, DOI: 10.15553/c2014v692a9, http://zenodo.org/record/5768228

Benstonea misimaensis (St. John ex Stone 1978: 54) Callm., Buerki & A.P. Keim, comb. nov. Basionym: Pandanus misimaensis St. John ex Stone (1978: 54). Type:— PAPUA NEW GUINEA. Milne Bay Province: Misima Island, Mt Sisa, N slopes, 300 m, 19 July 1956, fr., Brass 27401 (holotype, L [L0050582] image seen; isotype, A [A00020059] image seen, LAE, S [S-G-406] image seen) Notes: — Benstonea misimaensis is the smallest species in the ‘Dimissistyli’ group. The species is almost acaulescent, with the stem only attaining approximately 20 cm in height and grows in understory forest. This habit and habitat are similar to those of Benstonea ornithocephala (Stone 1978: 64) Callmander & Buerki in Callmander & al. (2013: 59). The two species are nevertheless very easily to distinguish, B. ornithocephala having a terminal syncarp and the upper stylar part of drupe not deciduous at maturity is not a member of the ‘Dimissistyli’ group. Benstonea misimaensis is endemic to the Island of Misima, 120 km to the east of mainland Papua New Guinea, in the Louisiade Archipelago, whereas B. ornithocephala is endemic to Bird’s Head Peninsula in West Papua Province (Indonesia). Both species are only known from the type collection.

Benstonea jacobsii (Stone 1984: 210) Callm., Buerki & A.P. Keim, comb. nov. Basionym: Pandanus jacobsii Stone (1984: 210). Type:— PAPUA NEW GUINEA. Hela Province: Near Waro airstrip, 20 km SSW of Kutubu, 6°31’S 143°10’E, 500–600 m, 15 October 1973, Jacobs 9281 (holotype, L [L0050543, L0050544] images seen). Notes: — Benstonea jacobsii was described by Stone (1984) in the genus Pandanus based on the small dimensions of its leaves (220 x 5 cm) and fruits (c. 10–11 cm in diam.) and the relatively smooth coarsely verruculose pileus. It is a small treelet (2–3 m) growing on limestone in the southeastern region of Papua New Guinea. Benstonea jacobsii can be easily distinguished from B. stenocarpa, by its habit (tall slender tree up to 10 m) and morphology (long and broad leaves, 400 x 11 cm) (Fig. 1F). Benstonea jacobsii is only known by the type specimen and is apparently restricted to limestone., Published as part of Callmander, Martin W., Buerki, Sven, Keim, Ary P. & Phillipson, Peter B., 2014, Notes on Benstonea (Pandanaceae) from the islands of Halmahera, New Guinea and Sulawesi, pp. 161-165 in Phytotaxa 175 (3) on page 163, DOI: 10.11646/phytotaxa.175.3.6, http://zenodo.org/record/5143877, {"references":["Stone, B. C. (1984) Some new and critical Pandanus species of subgenus Acrostigma, I. Supplement to Revisio Pandanacearum. Gardens' Bulletin Singapore 36: 205 - 212."]}

Benstonea misimaensis (St. John ex Stone 1978: 54) Callm., Buerki & A.P. Keim, comb. nov. Basionym: Pandanus misimaensis St. John ex Stone (1978: 54). Type:— PAPUA NEW GUINEA. Milne Bay Province: Misima Island, Mt Sisa, N slopes, 300 m, 19 July 1956, fr., Brass 27401 (holotype, L [L0050582] image seen; isotype, A [A00020059] image seen, LAE, S [S-G-406] image seen) Notes: — Benstonea misimaensis is the smallest species in the ‘Dimissistyli’ group. The species is almost acaulescent, with the stem only attaining approximately 20 cm in height and grows in understory forest. This habit and habitat are similar to those of Benstonea ornithocephala (Stone 1978: 64) Callmander & Buerki in Callmander & al. (2013: 59). The two species are nevertheless very easily to distinguish, B. ornithocephala having a terminal syncarp and the upper stylar part of drupe not deciduous at maturity is not a member of the ‘Dimissistyli’ group. Benstonea misimaensis is endemic to the Island of Misima, 120 km to the east of mainland Papua New Guinea, in the Louisiade Archipelago, whereas B. ornithocephala is endemic to Bird’s Head Peninsula in West Papua Province (Indonesia). Both species are only known from the type collection., Published as part of Callmander, Martin W., Buerki, Sven, Keim, Ary P. & Phillipson, Peter B., 2014, Notes on Benstonea (Pandanaceae) from the islands of Halmahera, New Guinea and Sulawesi, pp. 161-165 in Phytotaxa 175 (3) on page 163, DOI: 10.11646/phytotaxa.175.3.6, http://zenodo.org/record/5143877, {"references":["Stone, B. C. (1978) Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesaina precursores. Federation Museums Journal, new series 23: 1 - 74.","Callmander, M. W., Booth, T. J., Beentje, H. & Buerki, S. (2013) Update on the systematics of Benstonea (Pandanaceae): When a visionary taxonomist foresees phylogenetic relationships. Phytotaxa 112: 57 - 60. http: // dx. doi. org / 10.11646 / phytotaxa. 112.2.4"]}

Benstonea celebica (Warburg 1900: 80) Callmander & Buerki in Callmander & al. (2012: 332). Basionym: Pandanus celebicus Warburg (1900: 80). Type:��� INDONESIA. Sulawesi: s.d., Warburg s.n. (holotype, B [B100279969] image seen; isotype, FI [FI 003942]!). Epitype (designated here):��� INDONESIA. Central Sulawesi Province: Sulawesi, Mt Roroka Timbu, W slope, 80 km SSE of Palu, 1��16���S, 120��18���E, c. 1050 m, 11.V.1979, Vogel 5287 (epitype, L [L0332713] image seen; isoepitype, KLU-36404 [2 sheets] images seen, PH [PH 00018268]!). Syn.: Pandanus gladiator Stone (1984: 209). Type:��� INDONESIA. Central Sulawesi Province: Sulawesi, Mt Roroka Timbu, W slope, 80 km SSE of Palu, 1��16���S, 120��18���E, c. 1050 m, 11.V.1979, Vogel 5287 (holotype, L [L0332713] image seen; isotype, KLU-36404 [2 sheets] images seen, PH [PH 00018268]!). Notes:��� Benstonea celebica was regarded as an enigmatic species before its rediscovery by AK in 2005 (Keim 558) in Sulawesi at Pontolotua Hill during a survey of the Lore-Lindu National Park (Keim, 2005). The delimitation of this taxon has remained challenging for a century due to the very fragmentary material of the type (Warburg s.n.). The fruiting material deposited at B was apparently destroyed during World War II, but a photograph exists at FI. Stone (1978) has even suggested that B. celebica and B. pachyphylla (Merrill 1922: 154) Callmander & al. (2012: 337) may be conspecific. Benstonea celebica is in reality very different from B. pachyphylla, which is a slender acaulescent shrub bearing a small syncarp with a small infrafoliar near-ground solitary syncarp (see Keim & al. 2011: 52, Fig. 12; Callmander & al. 2012: 331, Fig. 4D). Benstonea celebica is a tree up to 5 m with a large spicate pendulous infructescence comprising up to 10 subspherical syncarps (c. 10���11 cm in diam) (Fig. 1G). In the absence of adequate material of B. celebica and apparently unaware of the presence of the photograph at FI, Stone (1984) was evidently oblivious to its highly distinctive morphology, and he described Pandanus gladiator Stone (1984: 209) based on a specimen collected from the same island, Sulawesi, Vogel 5287. The latter species was erroneously considered to be a synonym of Benstonea stenocarpa by Callmander & al. (2012) and this error is corrected here. Original material of B. celebica comprises only Warburg���s type gathering from Sulawesi, which is highly fragmentary - consisting now of only of two incomplete leaf portions (at B and FI). The only known syncarp of this collection is believed to have been destroyed (at B), although fortunately a photograph is conserved at FI. Vogel 5287 (the type of Pandanus gladiator) is an excellent fertile collection from Central Sulawesi with duplicates in L, KLU and PH, each with leaves and representative portions of the infructescence. We consider this specimen to represent the same species as the type of B. celebica, and we therefore designate it as an epitype for the species. Benstonea celebica has also been collected recently elsewhere on Sulawesi: Central Sulawesi Province, Pontolotua Hill and Sedoa in the vicinity of the Lore-Lindu National Park (Keim, 2005; Keim & Rustiami, 2007) and in Northern Sulawesi Province, Ambang Hill in the vicinity of Boganinani Wartabone National Park and Bolaang Mongondow, Minahassa (Polosakan & al. 2009; Purwanto & al. 2009) and is considered to be endemic to the island. Additional specimens seen: ��� INDONESIA. Central Sulawesi Province: Sulawesi, Toro, Pono (Pontolotua) Hill, 900 m, 31 May 2005, Keim 558 (WAL); Poso, Sedoa, Siboala Hill, 11 September 2005, Keim 683 (BO). North Sulawesi Province: Sulawesi, Mount Ambang Nature Reserve, near Kotulidak River, 00��47���29���N, 124��22���39���E, 23 November 2009, Kinho 267 (WAL); Bolaang Mongondow, Kolangkangon, around Ambang mountain, Modayag, 00��44���45���N, 124��26���07���E, 1271 m, 27 March 2006, Purwanto & al. PSU42 (BO)., Published as part of Callmander, Martin W., Buerki, Sven, Keim, Ary P. & Phillipson, Peter B., 2014, Notes on Benstonea (Pandanaceae) from the islands of Halmahera, New Guinea and Sulawesi, pp. 161-165 in Phytotaxa 175 (3) on pages 162-163, DOI: 10.11646/phytotaxa.175.3.6, http://zenodo.org/record/5143877, {"references":["Warburg, O. (1900) Pandanaceae. In: Engler, A. (Ed.) Das Planzenreich 3 (IV, 9). Leipzig, Engelmann, Germany, 97 pp.","Callmander, M. W., Lowry II, P. P., Forest, F., Devey, D. S., Beentje, H. & Buerki, S. (2012) Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species. Candollea 67: 323 - 345.","Stone, B. C. (1984) Some new and critical Pandanus species of subgenus Acrostigma, I. Supplement to Revisio Pandanacearum. Gardens' Bulletin Singapore 36: 205 - 212.","Keim, A. P. (2005) A joint Herbarium Bogoriense & HerbariumWallacea exploration in the Lore Lindu National Park, Central Celebes (including Mount Nokilalaki & Adjacent areas): Pandanaceae. Technical report, LIPI, Bogor, 18 pp.","Stone, B. C. (1978) Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesaina precursores. Federation Museums Journal, new series 23: 1 - 74.","Merrill, E. D. (1922) New or noteworthy Bornean plants, part 1. Journal of the Straits Branch of the Royal Asiatic Society 85: 151 - 201.","Keim, A. P., Rugayah & Rustiami, H. (2011) The Pandanaceae of the Bukit Baka Bukit Raya National Park and adjacent areas, West and Central Kalimantan, Indonesia, with notes an their nomenclature and the rediscovery of Pandanus aristatus and several new records. Garden's Bulletin Singapore 63: 31 - 62.","Keim, A. P. & Rustiami, H. (2007) Kenakeragaman suku Pandanaceae di pegunungan sekitar desa Sedoa, kawasan Taman Nasional Lore Lindu Kabupaten Poso-Propinsi Sulawesi Tengah. Berita Biologi 8: 375 - 389.","Polosakan, R., Susiarti, S. & Purwanto, Y. (2009) Habitat Pandanus di Gunung Ambang Taman Nasional Boganinani Wartabone dan pemanfaatan oleh masyarakat sekitarnya. In: Nuryanto, A. & Budisantoso, I. (Eds.) Prosiding Seminar Nasional Biologi Peran Biosistematika dalam Pengelolaan Sumberdaya Hayati Indonesia. General Soedirman University, Purwokerto, pp. 748 - 753.","Purwanto, Y., Polosakan, R. & Susiarti, S. (2009) Studi etnobiologi Pandanaceae masyarakat Minahassa & Bolaang Mongondow. Prosiding Seminar Nasional Etnobotani 4: 151 - 167."]}

Benstonea verruculosa (Backer ex Stone 1978: 55) Callm., Buerki & Phillipson, comb. nov. Basionym: Pandanus verruculosus Backer ex Stone (1978: 55). Type:— INDONESIA. North Maluku Province: Halmahera, Galileo, Sao Tobaroe, Beguin 1920 (holotype, BO [2 sheets]!, isotype, L [L0050725] image seen). Notes: — Benstonea verruculosa was described based on a fragmentary collection from Halmahera dating back to 1921 (Beguin 1920). Its overall morphology bears a strong resemblance to B. stenocarpa but its recent rediscovery on the island and collection of more adequate material (Callmander & Lasut 1151) provides a clarification of its delimitation and ample justification for its recognition at species level. Benstonea verruculosa is characterized by larger and broader leaves (c. 450 x 14 cm) (400 x 11 cm in B. stenocarapa), but more significant is the distinct morphology of its drupes. The style of B. verruculosa are short (Fig. 1A–G), and not elongated and gradually tapering as in B. stenocarpa. The stigmatic groove is also accordingly much shorter in B. verruculosa. Furthermore, the pileus of B. verruculosa is conspicuously verrucate contrasting with the rather smooth pileus (rarely roughly granular) of B. stenocarpa. The type of B. verruculosa is fragmentary, consisting of only leaf portions and separate drupes. Callmander & Lasut 1151 is the first complete collection of B. verruculosa, a species endemic to Halmahera where it grows along streams. Additional specimens seen:— INDONESIA. North Maluku Province: Halmahera, Lolea Poeloe Rao, 40 m, 12 October 1921, Beguin 1816 (BO, L); Soa Tobaroe, 60 m, 2 June 1922, Beguin 1933 (BO); Soa Tobaroe, 60 m, 13 August 1922, Beguin 2119 (BO); Soa Tobaroe, 60 m, 13 August 1922, Beguin 2120 (BO, L); Central Halmahera, Doromesmesan, Loy Poloy, 00°30’29”N, 127°54’21”E, 15 m, 9 February 2013, Callmander & Lasut 1151 (BO, G, L, MO)., Published as part of Callmander, Martin W., Buerki, Sven, Keim, Ary P. & Phillipson, Peter B., 2014, Notes on Benstonea (Pandanaceae) from the islands of Halmahera, New Guinea and Sulawesi, pp. 161-165 in Phytotaxa 175 (3) on page 163, DOI: 10.11646/phytotaxa.175.3.6, http://zenodo.org/record/5143877, {"references":["Stone, B. C. (1978) Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesaina precursores. Federation Museums Journal, new series 23: 1 - 74."]}

Pandanus Parkinson sect. Bernardia B.C.Stone ex Callm., sect. nov. Pandanus Parkinson sect. Bernardia B.C.Stone, Adansonia , s��r. 2, 12: 418 (1972), nom. inval. TYPUS. ��� Pandanus bernardii H.St.John ex Callm., sp. nov. DIAGNOSIS Treelet to 2-4 m tall; infructescence, a solitary, terminal subglobose syncarp; drupes obovoid truncate, slightly compressed, with deep and close ridges longitudinally; carpels 6-15-locular; pileus apex truncate with an apical concavity, flat to concave, gathering the stigmas; stigmas generally deltoid but sometimes deeply bifid, topped by an 1-3 mm long, appressed style often overlapping dorso-ventrally in 2(-3) rows., Published as part of Callmander, Martin W., 2014, Validating a striking new species endemic from New Caledonia: Pandanus bernardii H. St. John ex Callm., sp. nov. (Pandanaceae) and its monospecific section: Pandanus Parkinson sect. Bernardia B. C. Stone ex Callm., sect. nov., pp. 45-51 in Adansonia (3) (3) 36 (1) on page 46, DOI: 10.5252/a2014n1a4, http://zenodo.org/record/5208398

Callmander, Martin W. (2014): Validating a striking new species endemic from New Caledonia: Pandanus bernardii H.St.John ex Callm., sp. nov. (Pandanaceae) and its monospecific section: Pandanus Parkinson sect. Bernardia B.C.Stone ex Callm., sect. nov. Adansonia (3) 36 (1): 45-51, DOI: 10.5252/a2014n1a4, URL: http://dx.doi.org/10.5252/a2014n1a4

Pandanus bernardii H.St.John ex Callm., sp. nov. (Fig. 2) Pandanus bernardii H.St.John ex B.C.Stone, Adansonia, s��r. 2, 12: 418 (1972); Botanical Magazine of Tokyo 95: 230 (1982); Revision of the genus Pandanus Stickman. Part 61. Pandanus of New Caledonia: 11 (1989), nom. nud. Haec species inter congeneros novacaledonicos syncarpio subgloboso pallide viridi 12-15 �� 11-13 cm, drupa obovoidea truncata leviter compressa longitudinaliter profunde crebiterque porcata atque stigmatibus 6 ad 15 plerumque deltoideis sed interdum profunde bifidis secus stylum adpressum elevatis (1-3 mm) saepe in seriebus 2 (vel 3) dorsoventraliter imbricatis distinguitur. TYPUS. ��� New Caledonia. Grande Terre, Rivi��re Bleue, in regione submontana, [22��05���30������S, 166��41���30������E], 200 m, 4.VII.1965, fr., Bernardi 9431bis (holo-, G [G00305792]!; iso-, BISH!, MO!, NOU!, P [P00700914]!, PH!). PARATYPI. ��� New Caledonia. Grande Terre. Mandj��lia, 20��24���S, 164��32���E, 610 m, 4.I.1978, fr., Bamps 6140 (BM, NOU [NOU048601]). ��� For��t Demazures, vall��e La Coul��e, [22��10���44������S, 166��36���59������E], 10.III.2005, fr., Barrab�� 295 (NOU [NOU004603]). ��� Mois de Mai, rivi��re blanche sup., [22��08���59������S, 166��36���34������E], 300 m, 25.VI.1951, fr., Baumann-Bodenheim & Guillaumin 14308 (G, BISH, P [P02077319]). ��� Pic du Pin, versant SE, [22��14���34������S, 166��49���31������E], 450 m, 14.II.2008, fr., Dagostini 1570 (NOU [NOU031140, NOU031141, NOU050577]). ��� For��t des ��oliennes, en contrebas du col de Prony, [22��18���S, 166��45���E], 190 m, 1.XI.2007, fr., Fambart-Tinel (leg. Barriera) 156 (NOU [NOU024184, NOU050588]). ��� En dessus de la cabane des ��lectriques, [22��10���11������S, 166��39���32������E], 240 m, 3.IX.1958, fr., H��rlimann 3422 (BISH [mixed with P. pancheri (Brongn.) Solms], G). ��� Route de Prony, [22��18���30������S, 166��46���E], 10.VII.1984, fr., Jaffr�� 2509 (NOU [NOU050553, NOU048600], P [P02077371], PH). ��� Haute Rivi��re Bleue, [22��05���26������S, 166��41���30������E], 150-200 m, 27.IX.1967, fr., MacKee 17590 (P [P02077372, carpo.]). ��� Haute Diahot, for��t Tend��, [20��24���37������S, 164��31���17������E], 500 m, 30.IX.1968, fr., MacKee 19677 (G [carpo.], FI [carpo.], NOU [NOU048593], P [P02077373, carpo.]). ��� Pou��bo, cr��te entre Mandj��lia et Salandan��, [20��23���12������S, 164��31���17������E], 600 m, 15.XII.1969, fr., MacKee 21287 (G [carpo.], MO [carpo.], NOU [NOU048590], P [P02077387, carpo.]). ��� ibid. loc., st. fl., MacKee 21289 (G, NOU [NOU048589], P [P02077369]). ��� Haute N��aoua, Ouen Sieu, [21��23���12������S, 165��32���13������E], 500 m, 12.II.1970, st. fl., MacKee 21607 (G, NOU [NOU048591, NOU048592], P [P02077370, carpo.]). ��� Pou��bo, Ouangati, [20��26���25������S, 164��34���20������E], 800 m, 26.V.1971, fr., MacKee 23791 (G, FI [carpo.], MO [carpo.], NOU [NOU048585, NOU048586], P [P02077368, carpo.]). ��� Cr��te au S Baie Ouinn��, [21��59���51������S, 166��40���15������E], 400-600 m, fr., 10.XI.1973, MacKee 27774 (G, FI, MO, NOU [NOU048587], P [P02139394]). ��� For��t Faux Bon Secours, [22��10���02������S, 166��41���33������E], 300 m, 13.XII.1979, fr., MacKee 37677 (G, NOU [NOU048595, NOU050551], P [P02077385, carpo.], PH). ��� Haute Ku��bini, [22��13���53������S, 166��57���48������E], 400 m, 29.X.1981, fr., MacKee 39839 (NOU [NOU050555], P [P02082014, carpo.], PH [2 sheets]). ��� Mt Pani��, c. 20 air-km NW of Hiengh��ne, [22��13���53������S, 166��57���48������E], 500 m, fr., 3.III.1981, McPherson 3691 (MO, NOU [NOU048599], P [P02077383], PH). ��� R��serve du Pic du Grand Kaori, [22��17���31������S, 166��53���37������E], 30.VIII.2007, fr., Munzinger 4503 (NOU [NOU029159]). ��� Yat�� Barrage, 21.VIII.2008, fr., Munzinger 5082 (NOU [NOU050788]). ��� For��t cach��e, [22��11���20������S, 166��47���10 ������E], fr., Munzinger s.n. (NOU [NOU050597]). ��� Vall��e de la Pourina, 13.XI.2008, 22��02���58������S, 166��43���23������E, fr., Pillon 1337 (NOU [NOU049504]). ��� Hills above La Riv. Bleue, [22��05���26������S, 166��41���30������E], 550 m, 30.VIII.1971, fr., Stone 10547 (BISH, G, K, NOU [NOU048580], PH). ��� Riv. Bleue, route d���acc��s �� la haute Pourina, [22��05���26������S, 166��41���30������E], 400 m, 25.XI.1982, st. fl., Suprin 2175 (NOU [NOU048596, NOU048597], PH). ��� Yat��, Rivi��re Bleue (apr��s la cabane), [22��06���46������S, 166��41���04������E], c. 250 m, 6.IX.1967, fr., Veillon 1339 (NOU [NOU048598, NOU050550], P [P02077381, P02077382, carpo.]). ��� Le Cresson, exploitation Frouin, [20��29���14������S, 164��18���17������E], 500 m, 28.XI.1967, fr., Veillon 1573 (NOU [NOU048584]). ��� Plaine des Lacs, 19.XII.1969, [22��16���13������S, 166��56���59������E], fr., Veillon 2082 (BISH, NOU [NOU048581, NOU048582, NOU048583]). ��� Plaine des Lacs, [22��16���13������S, 166��56���59������E], 19.XII.1969, st. fl., Veillon 2082A (BISH). ��� La Rivi��re Bleue, pr��s du Grand Kaori, [22��05���59������S, 166��40���21������E], X.1979, st. fl., Veillon 4212 (NOU [NOU048499], P [P02081988], PH). DISTRIBUTION AND ECOLOGY. ��� Pandanus bernardii H.St. John ex Callm., sp. nov. is known from lowland humid forests from 200-800 m in southern and northeastern New Caledonia (Fig. 1). It grows on schist and serpentine. ETYMOLOGY. ��� This species and the monospecific section it belongs to is named in honour of Luciano Bernardi (1920-2001), former curator at the ���Conservatoire et Jardin botaniques de la Ville de Gen��ve���. Bernardi, an ���infatigable explorateur���, is known for his numerous field trips around the world, especially in Africa, Asia, Madagascar, South America and the South Pacific (see Spichiger et al. 2002). Over tens of thousands of specimens, among which the type of the species described here, were gathered in the course of his expeditions. CONSERVATION STATUS. ��� With an EOO of 9 020 km2, an AOO of 69 km 2, and six subpopulations (calculations following Callmander et al. 2007), two of which are situated within the protected area network (For��t Cach��e, Montagne des Sources, Mt. Pani��, Pic du Grand Kaori, Pic du Pin, Rivi��re Bleue), Pandanus bernardii H.St. John ex Callm., sp. nov. is assigned a preliminary status of Vulnerable (VU B1ab[iii]+ B2ab[iii]) based on the IUCN Red List Categories and Criteria (IUCN 2012). DESCRIPTION Treelet to 3-4 m tall, stem 10-12 cm in diam., with few to several prop-roots to 20 cm high. Leaves linear-attenuate, 100-150 cm long, 3-5 cm wide, apex acute; thin coriaceous, green, shiny abaxially, pale green adaxially; longitudinal veins visible on both surfaces; marginal prickles borne (8-) 10-14cm above base to the apex, antrorse, 0.3-0.7 mm long in the proximal third, 1.5-4.0 mm apart, 0.2-0.5 mm long in the mid-third, similarly spaced, to 0.1-0.4mm long in the distal third, 0.5-2.0 mm apart, subappressed; midrib unarmed in the lower (1/3-)1/2, prickles> 0.3 mm long towards the apex, irregularly spaced but mostly 1-3 mm apart; apical ventral pleats usually with a few antrorse prickles, 0.3-0.6 mm long, mostly 2-5 mm apart, sometimes lacking; sheath 8-10 cm long, c. 3.5-5 cm wide at apex, 6-7 cm at base. Infructescence terminal, the solitary syncarp, subglobose, 12-15 cm long, 11-13 cm in diam., pale green; peduncle 18-22 cm long, 1.6 cm thick at apex, trigonous, with several foliaceous bracts, the lower ones about 75 cm long and decreasing in size towards the apex. Drupes c. 50-110, connate in the mature syncarp, c. (36-) 40-60 mm long, (18-) 30-40 mm wide, 15-22 mm thick, mostly (6-)7-15-locular, 4-7-angled, obovoid-truncate, slightly compressed, with deep and close ridges longitudinally; pileus apex flat to concave. Stigmas 6-15, c. 1.5-2.5 mm wide, generally deltoid but sometimes deeply bifid, topped by a 1-3 mm long appressed style, proximal or sometimes facing each other, often overlapping. Endocarp median, 10-12 mm long, distally subtruncate margins slightly ascending; walls 2-3 mm thick but with slight conic projections over the seeds, the shoulders small but ascending, acute, lateral walls 2-3 mm thick; proximal mesocarp fibrous fleshy, equally long or longer, distal mesocarp c. 15 mm long, dense, fibrous and medullate; seed locule ellipsoid, 6-7 mm long. Staminate inflorescence a raceme of spikes c. 36 cm long; peduncle soft, fleshy; lowest floral bract 34 cm long, 5 cm wide, lanceolate, cuneate, navicular, foliaceous, the margins and midrib finely serrulate, the uppermost bract 7.5 cm long, 2.8 cm wide, lanceolate, foliaceous like the others; spikes c. 7-8, 5-9(- 15 cm) long, 2-3 cm wide, composed of crowded ���stemanophore���; ���stemanophore��� with column 6-8 mm long, c. 2-3 mm wide, apically distended, umbrella like,peltate, to c. 6 mm wide, blackish when dry; stamens c. 80, subsessile, borne along upper part of ���stemanophores��� and (mostly) under the apical discoid surface, spreading from its rim; filament c. 0.2 mm long; anthers 1.5-1.6 mm long, 0.3 mm wide, oblong, apiculate, apiculus 0.2 mm long. REMARKS With its deflected stigmas and staminate flower characterized by a ���stemanophore���, a fascicle of stamens, expanded into a peltate, umbrella like surface (Fig. 2I), Pandanus bernardii H.St.John ex Callm., sp. nov. can be without any doubt placed in Pandanus subg. Lophostigma (Brongn.) H.St.John to whom all New Caledonian species belong except the widespread Pandanus tectorius (type of Pandanus subg. Pandanus). A recent molecular phylogenetic analysis including the new species described here confirms the monophyly of all New Caledonian species except the widespread P. tectorius (Buerki et al. 2012). Nevertheless, the new species has a unique combination of morphological characters and cannot be confused to any other known Pandanus species and therefore is placed on its own monospecific section following Stone���s infrageneric classification (1974). The new species can be recognized by its unlobed obovoid drupes, with numerous and parallel longitudinal deep ridges that runs from the base to the apex of each drupe (Fig. 2 B-E). The drupe apex is truncate and holds a concavity, where all the stigmas are gathered. The stigmas are generally deltoid but sometimes deeply bifid, topped by a 1-3 mm long appressed style, proximal or sometimes facing each other and somewhat overlapping (Fig. 2B)., Published as part of Callmander, Martin W., 2014, Validating a striking new species endemic from New Caledonia: Pandanus bernardii H. St. John ex Callm., sp. nov. (Pandanaceae) and its monospecific section: Pandanus Parkinson sect. Bernardia B. C. Stone ex Callm., sect. nov., pp. 45-51 in Adansonia (3) (3) 36 (1) on pages 47-50, DOI: 10.5252/a2014n1a4, http://zenodo.org/record/5208398, {"references":["SPICHIGER R., MAIULLARI F., WENGER M. & PERRET P. 2002. - Dr. Luciano Bernardi, felsineus (\" celui de Bologne \") (6.3.1920 - 1.12.2001). Candollea 56: 363 - 376.","CALLMANDER M. W., SCHATZ G. E., LOWRY II P. P., LAIVAO M. O., RAHARIMAMPIONONA J., ANDRIAM- BOLOLONERA S., RAMINOSOA T. & CONSIGLIO T. 2007. - Application of IUCN Red List criteria and assessment of Priority Areas for Plant Conservation in Madagascar: rare and threatened Pandanaceae indicate new sites in need of protection. Oryx 42: 168 - 176.","IUCN 2012. - IUCN Red List Categories and Criteria: Version 3.1. 2 nd Edition, IUCN Species Survival Commission, Gland & Cambridge, ii + 30 p.","ST. JOHN H 1982. - Revision of the genus Pandanus Stickman, Part 45. New Pandanus species (Pandanaceae) from New Caledonia. Botanical Magazine of Tokyo 95: 229 - 239.","BUERKI S., CALLMANDER M. W., DEVEY D. S., CHAPELL L., GALLAHER T., MUNZINGER J., HAEVERMANS T. & FOR- EST F. 2012. - Straightening out the screw-pines: a first step in understanding phylogenetic relationships within Pandanaceae. Taxon 61: 1010 - 1020."]}

Benstonea ornithocephala (B.C. Stone) Callm. & Buerki, comb. nov. Basionym: Pandanus ornithocephalus Stone (1978: 64). Type:— INDONESIA [NEW GUINEA]. West Papua Prov.: Vogelkop Peninsula [Kepala Burung], Skendi (N of Teminaboean), c. 100 m, 21 May 1958, fr., Versteegh BW 7474 (holotype, L [L0050607]!), Published as part of Callmander, Martin W. & Buerki, Sven, 2013, Corrections to Phytotaxa 112: Update on the systematics of Benstonea (Pandanaceae), pp. 59-60 in Phytotaxa 125 (1) on page 59, DOI: 10.11646/phytotaxa.125.1.9, http://zenodo.org/record/5085206, {"references":["Stone, B. C. (1978) Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesiana precursores. Federation Museums Journal, new series 23: 1 - 74."]}

Benstonea ornithocephala (B.C. Stone) Callm. & Buerki, comb. nov. Basionym: Pandanus ornithocephalus Stone (1978: 64). Type:— INDONESIA [NEW GUINEA]. West Papua Prov.: Vogelkop Peninsula [Kepala Burung], Skendi (N of Teminaboean), c. 100 m, 21 May 1958, fr., Versteegh BW 7474 (holotype, L [L0050607]!)

Pandanus reticulatus Vieill. in Ann. Sci. Nat., Bot. ser. 4, 16: 52. 1862 (Fig. 1, 3B, 4). Neotypus (designated here): NEW CALEDONIA: R��serve botanique de Nod��la, flan du massif du M�� Moya, 21��26���07���S 165��20���05���E, 500 m, 19.II.2011, fr., Callmander & al. 981 (P [P00700916]!; iso-, BISH!, BM!, FI!, G [G00368275]!, K!, MO!, NOU [NOU054165]!, P [P007 00915]!, PH!). Understory erect shrub to 3 m tall, with slender weak, sometimes leaning, stem, and a few branches; prop roots few, but up to 1 m long; leaf crowns dense. Leaves linear-attenuate, (50-)75-80(-90) cm long, 1.2-1.5 cm wide, attenuate or but slightly acuminate toward apex; subcoriacous in the proximal half part, chartaceous in the distal half, discolour, dark green to brown abaxially, pale green adaxially when dried; longitudinal veins visible on both surfaces; marginal prickels borne 2-2.5 cm above base to apex, antrorse, 1 mm long, 4-7 mm apart in the proximal third, ca. 1 mm, 2-5 mm apart in the mid third, 0.5-1 mm long, 1-2 mm apart in the distal third, more appressed towards the apex; midrib near base unarmed to beyond the middle, after with sparse, remote prickles similar to those adjacent to margins, ca. 0.5 mm long; apical ventral pleats with a few antrorse prickles, unequally dispersed, often mostly or only on one (usually the left) side, lacking on extreme tip. Infructescence solitary, terminal, the syncarp oblong-ovoid, ca. 2.5-3 �� 5-6 cm, on short erect, peduncle about 8 cm long, 4-5 mm thick. Drupes ca. 90-110, 10- 15 mm long, 5-10 mm wide, 7-10 mm wide, mostly l-celled, very rarely 2-celled, 5-6-angled, with smooth sides; pileus 3 mm long, crowned by the antrorsely excentric inclined stigma. Stigma unique, rounded reniform, 1-1.5 mm wide. Proximal mesocarp fibrous, 3-4.5 mm long, distal mesocarp fibrous, 2-4 mm long; endocarp sub-median or median, about 5 mm long, the walls ca. 1 mm thick. Seed locule 3 mm long. Staminate inflorescence to ca. 20 cm long; proximal bracts foliaceous-tipped; distal bract ca. 3 �� 15 cm and decrescent, margins and midrib finely ciliate, serrulate-prickly, navicularovate in outline; spikes about 5-6, short sub-cylindric, ca. 25- 50 mm long, terminal spike longer, ca. 7-8 cm, merged at base with adjacent lateral spike, composed of crowded phalanges of stamens. Stemanophore sub-peltate, the column, ca. 2 mm in length, with a deep inverse-conic depression in the distal part, the margins bearing the stamens radially. Stamens about 13-18 per stemanophore, the filament 0.75-1 mm long; a short filamentous pendiculus intervening between filament and anther, the pendiculus 0.1-0.2 mm long. Anther oblong, acuminate, ca. 1 mm long, with an apiculus ca. 0.35 mm long, dorsally set with raphidophorus cells, at apex subentire; connective dorsally and distally with a few raphidophorus cells. Distribution and ecology. ��� Pandanus reticulatus is known from lowland dense humid forest in northeastern and southern New Caledonia between 300-700 m on schist and serpentine (Fig. 1). Conservation status. ��� With an EOO of 6979 km 2, an AOO of 90 km 2, and 6 subpopulations, one of which is situated within the protected area network (Barrage Yat��, Haute Yat��, Rivi��re Bleue, Pic du Grand Kaori), P. reticulatus is assigned a preliminary status of Vulnerable (VU B1ab [iii]; B2ab [iii]) based on the IUCN Red List Categories and Criteria (IUCN, 2001). Notes. ��� Pandanus reticulatus was described by VIEILLARD (1864) very succinctly. This species was considered by MARTELLI (1914) and STONE (1972) to be synonym of P. decumbens, whereas ST. JOHN (1989: 25) treated it as a ���species dubia���. BRONGNIART (1875: 286) noted the close resemblance but underlined the morphological distinction by emphasising Vieillard���s description: ���drupe petite, serr��es, subtetragones, couron��es par le stigmate persistant, uniloculaire���, whereas P. decumbens holds larger drupes with several stigmas in one row. No type material has been located, nor was any referred to by MARTELLI (1914), St. JOHN (1989) or Stone (pers. comm.), therefore a neotype is required to be designated. No collections are known to exist that Vieillard might have seen. Nevertheless, the species has been collected many times in the last 65 years before our visit to New Caledonia in 2011, but only one has a mature syncarp (MacKee 42975). During our visit we were able to locate a large population of P. reticulatus in Nod��la, on the slopes of the M�� Moya massif. This collection (Callmander & al. 981), which is accompanied by extensive field information and photographs and comprises multiple duplicates, is chosen as the neotype. The species is now described in detail above for the first time based on this collection. The species is unique among the New Caledonian taxa in having a unicarpellate drupe with a reniform stigma (Fig. 3B, 4). It can be easily recognized in the field by its slender habit usually with prop roots up to 1 m tall, its narrow leaves (ca. 12-15 mm wide) and tiny oblong ovoid syncarp (ca. 2.5 �� 5.5 cm) with unicarpellate drupes (Fig. 3B). Additional material examined. ��� NEW CALEDONIA : Riv. Blanche, Moi de Mai, [22��07���19���S 166��38���34���E], 300 m, 23.VI.1951, st., Baumann-Bodenheim 14099 (P [P02077341]); Dawenia, versant SW du Mont Colnett, 20��32���25���S 164��41���12���E, 575 m, 14.XI.2010, fr., Callmander & al. 877 (G, NOU, P [P00700842]); R��serve botanique de la Fausse Yat��, 22��11���05������S 166��54���14������E, 300-500 m, 9.X.2007, st. fl., Dagostini 1463 (NOU [NOU031137]); Riv. Bleue, [22��06���11���S 166��37���60���E], 200 m, 22.II.1951, st., Guillaumin & Baumann-Bodenheim 10906 (P [P02077341]); Yat��, [20��09���21���S 166��53���48���E], 300 m, 29.XII.1985, fr., MacKee 42975 (G, NOU [NOU006188], P [P02139400], PH [PH00072777, PH00072778, PH00072779]); Haute Rivi��re des Pirogues: For��t Faux Bon Secours, [22��10���02���S 166��41��� 33���E], 300 m, 12.I.1986, st., MacKee 42995 (NOU [NOU048442], P [P02139389], PH [PH00072771]); Pic du Grand Kaori, 22��17���19���S 166��53���48���E, 235 m, 9.X.2009, fr., Munzinger & al. 5978 (NOU); Hills above Riv. Bleue, [22��06���11���S 166��37���60���E], 27.VIII.1971, st., Stone 10551 (PH [PH00072773]); Ou��goa, bassin SW de la Tend��, [20��25��� 22���S 164��29���39���E], 22.VII.1973, fr., Veillon 2990 (NOU [NOU048440, NOU048441], PH [PH00072770]); Riv. Bleue, [22��011���S 166��37���60���E], 13.VI.1985, fr., Veillon 5910 (NOU [NOU048446], PH [PH00072775]); Riv. Bleue, [22��06���11���S 166��37��� 60���E], 200-250 m, 29.I.1986, fr., Veillon 5948 (NOU [NOU048447, NOU048448], PH [PH0072772]); Mt Remba��, [21��34���47���S 165��49��� 59���E], ca. 670 m, 17.XI.1989, st. fl., Veillon 7227 (NOU [NOU04 8439], P [P02139405])., Published as part of Callmander, Martin W. & Buerki, Sven, 2013, Notes on New Caledonian Pandanaceae: identity and typifications of Pandanus decumbens (Brongn.) Solms and Pandanus reticulatus Vieill., with the new species Pandanus letocartiorum Callm. & Buerki, pp. 51-60 in Candollea 68 (1) on pages 56-57, DOI: 10.15553/c2013v681a4, http://zenodo.org/record/5736361, {"references":["IUCN (2001). IUCN Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission, IUCN.","MARTELLI, U. (1914). Le specie e varieta nuove di \" Pandanus \" menzionate nella enumerazione delle Pandanaceae. Webbia 4: 399 - 435","STONE, B. C. (1972). On the genus Pandanus (Pandanaceae) in New Caledonia. Adansonia ser. 2, 12: 409 - 420.","ST. JOHN, H. (1989) Revision of the genus Pandanus Stickman, Part 61. Pandanus of New Caledonia. Harold St John, Honolulu.","BRONGNIART, A. (1875). Observations sur les Pandanees de la Nouvelle-Caledonie. Ann. Sci. Nat., Bot. ser. 6, 7: 262 - 293."]}

Pandanus decumbens (Brongn.) Solms in Linnaea 42: 47. 1878 (Fig. 1, 2, 3A). �� Barrotia decumbens Brongn. in Ann. Sci. Nat., Bot. ser. 6, 7: 285. 1875. Lectotypus: NEW CALEDONIA : Iles des Pins, sur les pentes du Pic [N���ga], [23��20���42������S 167��27���25������E], fr., Pancher 342 (P [P02077365]!; iso-: FI!, P [P02077364, P02077366, P02077367, P02139406]!) (lectotypified by ST. JOHN, 1989: 24). = Pandanus neocaledonicus Martelli in Webbia 4: 416. 1914. Lectotypus (designated here): N EW CALEDONIA : s.d., fr., s.coll. (FI!; iso-: P [P 00700817]!). = Pandanus schlechteri Warb. in Bot. Jahrb. Syst. 39: 17. 1906. Typus: NEW CALEDONIA : S��d-Bezirk. Auf den Bergen bei Yaouh��, [22��11���45������S 166��29���54������E], 400 m, X.1902, fr., Schlechter 15053 (holo-: B [B100 352291, image seen]!; iso-: FI!, P [P00649506]!). ��� Pandanus reticulatus sensu Martelli in Webbia 4: 29, 62, 94. 1913; 4: tab. 2, fig. 7-9. 1914. Slender-stemmed shrub, the stem usually decumbent, branched or not, often quite short but sometimes elongated, up to 4 m tall, with rather smooth light brown bark and a few distant blunt prickles; few prop roots, little developed, to ca. 20 cm long. Leaves linear-attenuate, mostly 100-135(-160) cm long, (15-)20-25(-50) mm wide, apex attenuate, sometimes sub-flagellate; thin sub-coriaceous to chartaceous, dark green and shiny axially, paler green and shiny adaxially; longitudinal veins visible on both surfaces; marginal prickles borne 7-8 cm above base to apex, antrorse, 0.5-1 mm long in the lower third, 2-8 mm apart, 0.4-0.6 mm long in the mid-third, 2-4 mm apart, to 0.2-0.4 mm long in the distal third, 1-2 mm apart, subappressed; midrib unarmed in the lower half, prickles> 0.2 mm long towards the apex, irregularly spaced but mostly 2-10 mm apart; apical ventral pleats with antrorse prickles,> 0.2 mm mm long, mostly 1-6 mm apart; sheath 5-7 cm long, ca. 2-3 cm wide at apex, 4-5 cm at base. Infructescence terminal, the solitary syncarp sub-cylindric, (3-) 7-15 cm long, (3-) 5-8 cm wide, peduncle ca. 8-15(-35) cm long, ca. 2 cm thick at apex, bearing foliaceous bracts, the proximal ones ca. 60-80 cm long, the distal one 12-14 cm long, crowded at peduncle apex and often covering the syncarp. Drupes numerous, 100-180, ca. 15-25 mm long, 10-20 mm wide, 4-10 mm thick, mostly (1-)3-4(-5)-celled, (4-)6-angled, obovoid, at apex truncate or shallowly concave, upper third free with smooth, dull, gently incurved faces. Stigma (1-)3-4(-5), ca. 1-2 mm wide, slightly heightened (ca. 2 mm) on an appressed style, forming one row, deflected toward the cephalium apex. Endocarp median, ca. 10 mm long, concave at apex, lateral walls 2-4 mm thick; proximal mesocarp 8-10 mm long, fibrous-fleshy, distal mesocarp 6-10 mm long, sparsely fibrous, chambered, seed locule ca. 4 �� 7 mm, ellipsoid. Staminate inflorescence a raceme of spikes, ca. 13-23 cm long; peduncle fleshy; bracts ca. 10, distal ones foliaceous, proximal ones caniculate, the distal bract ca. 2 �� 10 cm; spikes (4-)5, mostly 2-4 cm long, up to 15 mm diam., short cylindric, composed of crowded phalanges of stamens (stemanophore); its a column 4-6 mm long, apically distended, umbrella like, peltate, to ca. 4-5 mm wide, blackish when dry; stamens ca. 50-70, borne along upper part of the stemanophore and (mostly) under the apical discoid surface; filaments ca. 0.5 mm long; anthers 1.5- 2.4 mm long, to 0.5 mm wide, pale yellow, apiculate; apiculus 0.2 mm long. Distribution and ecology. ��� Pandanus decumbens is mostly found at lower to mid elevations (0-400 m). It grows on both serpentine and calcareous substrates, usually in rocky sites. The species is widespread along the west coast from Iles des Pins to Koumac usually in Maquis or dry forests and often along creeks (Fig. 1). Conservation status. ��� With an EOO of 15 642 km 2, an AOO of 324 km 2, and 18 subpopulations, three of which are situated within the protected area network (Barrage Yat��, Mt. Pani��, Vall��e de la Thy), P. decumbens is assigned a preliminary status of Least Concern (LC) based on the IUCN Red List Categories and Criteria (IUCN, 2001). Notes. ��� Pandanus decumbens was described by BRONGNIART (1875) under the basionym Barrotia decumbens Brongn., it was based on two syntypes: Balansa s.n. and Pancher 342. Balansa s.n. has not been located either at FI or at P. Pancher 342 is a complete collection comprising leaves, young and mature syncarps and drupes at P and separate drupes at FI. ST. JOHN (1989: 24) therefore designated this collection as the lectotype. The sheet of Pancher 342 at P [P020 77365] bears a label with ���n��8��� annotation (see ���sonneratphoto.mnhn.fr/ 2010/12/03 /2/P02077365.jpg��� for scanned specimen), as well as an original Pancher label with ���342��� clearly indicated. St. John wrongly cited the collection as Pancher 8 when he lectotypified the species. Note that St John actually cited the sheet as the holotype, but this is to be considered as an effective lectotypification since it was published before 2001 (see MCNEILL & al., 2012 [art. 9.23]). MARTELLI (1914) misinterpreted Pandanus decumbens and considered this species to be a synonym of P. reticulatus along with P. schlechteri Warb. He consequently described a new species, P.neocaledonicus Martelli, to accommodate a specimen (s.coll.) deposited at the Caen Museum. This material is believed to have been destroyed during World War II, but fragments of the collection have been found at FI and P. The FI sheet is designated as lectotype because it is in better condition and less fragmentary than the sheet at P. This collection clearly belongs to P. decumbens with its stigma slightly raised (ca. 2 mm) on an appressed style, arranged in one row and deflected toward the cephalium apex (Fig. 2, 3A). Pandanus neocaledonicus is therefore considered as a synonym of P. decumbens along with P. schlechteri, the latter already being treated in this way by ST. JOHN (1989). Additional material examined. ��� NEW CALEDONIA : Pic N���ga, [23��20���42������S 167��27���25������E], 29.V.1951, fr., Bauman-Bodenheim 13696 (K, P [P02077342], Z); Pic N���ga, creek sur les pentes SW, [23��20���42������S 167��27���25������E], 30.V.1951, st., Bauman-Bodenheim 13801 (P [P 02139407]); Ourou��, [21��34���59������S 166��11���32������E], 26.VI.1951, st., Bauman-Bodenheim 13696 (P [P 02077340]); Poum, 20��14���08������S 164��01���23������E, 13.IV.2006, fr., Barrab�� 356 (NOU [NOU 014166], P [P 02075145]); N. of Dumb��a valley, [22��08���15������S 166��29���05������E], 150 m, 7.VII.1947, fr., Buchholz 1178 (P [P 021 39409]); Mont Dzumac, 22��05���59���S 166��27���02���E, 572 m, 23.I.2010, fr., Callmander & al. 827 (G, NOU, P [P 00700824]); Sur la route de Gouapin, N��t��a, 21��14���32���S 165��14���50���E, 15 m, 16.II.2011, st. fl., Callmander & al. 976 (G, MO, NOU); Sur la route de Gouapin, N��t��a, 21��14���41���S 165��16���25���E, fr., Callmander & al. 977 (P [P 007 00859]); Massif du Boulinda, en dessous de la mine Saint-Louis, 21��18���48���S 165��06���12���E, 320 m, 18.II.2011, fr., Callmander & al. 978 (BISH, G, MO, K); Massif du Boulinda, en dessous de la mine Saint-Louis, 21��18���44���S 165��06���11���E, 365 m, fr., Callmander & al. 979 (MO); Massif de Poum, plaine c��t�� E, 20��15���37������S 164��02���17������E, 8.V.2008, y. fr., Dagostini 1603 (NOU [NOU 049031]); Iles des Pins, sur les pentes du Pic, [22��39���22������S 167��27���39������E], 1862, fr., Deplanche 143 (P [P 02077336]); Iles des Pins, 1874-1876, st., Germain s.n. (P [P 02077337, P 02077338, P 02077339]); Mt. Mou, [22��04���35������S 166��21���08������E], 15.X.1950, st., Guillaumin & Bauman- Bodenheim 6898 (P [P 02077335]); Mt. Mou, 300 m, [22��04���35������S 166��21���08������E], 15.X.1950, fr., Guillaumin & Bauman-Bodenheim 6953 (G, P [P 02139401], Z); Pic Tiaou��, 200-400 m, 21��10���46������S 165��01���24������E, 7.XI.2006, Hequet (Leg. Butin) 3661 (NOU [NOU 01 7072]); Ou��goa, St Andr�� Yallin, ca. 100 m, [20��19���47������S 164��26��� 29������E], 24.VII.1968, fr., Jaffr�� 60 (BISH); Dumb��a N. au bas du Col., rive droite, [22��08���15������S 166��29���05������E], 6.II.1970, st. fl., Jaffr�� 357 (BISH, NOU [NOU 048525, NOU 048526]); Dumb��a N. au bas du Col., rive droite, fr., Jaffr�� 358 (BISH, NOU [NOU048528]); vall��e de la Thy, [22��10���53������S 166��31���58������E], 3.VI.1973, fr., Jaffr�� 1128 (NOU [NOU048535]); Iles des Pins, base du Pic N���Ga, [22��39���18������S 167��27���25������E], 10.VIII.1975, fr., Jaffr�� 1443 (PH [PH 0072764]); Thy vall��e, [22��10���53������S 167��28���02������E], 50-100 m, 8.I.1956, st., MacKee 3748 (BISH, K, P [P 02077362]); Dumb��a Valley, [22��08���15������S 166��29���05������E], 100-200 m, 5.II.1956, fr., MacKee 3980 (BISH, K, P [P 02077361]); Dumb��a Valley below battage, [22��08���15������S 166��29���05������E], 100 m, 17.III.1956, y. fr., MacKee 4100 (BISH, K, P [carpo]); 7 km from Kone on road to Voh, [21��01���59������S 164��47��� 44������E], 10 m, 1.IV.1959, fr., MacKee 4309 (P [carpo]); Iles des Pins, slopes of Pic N���ga, [22��39���408������S 167��27���41������E], 100-200 m, 5.VII.1956, st., MacKee 5051 (P [P 02077353]); Ourou�� (embouchure de la Dothio), [21��34���59������S 166��11���32������E], 10-150 m, 20.X.1965, fr., MacKee 13633 (P [P 02077352, carpo]); Vall��e Koumac, [20��31���30������S 164��22���58������E], 21.VI.1967, fr., MacKee 16968 (BISH, FI [carpo], NOU [NOU 048532], P [P 02077351, carpo]); Voh, Base SE, Tahaf��, [20��57���44������S 165��15���35������E], 7.VII.1970, y. fr., MacKee 22166 (BISH, NOU [NOU 048513], P [P 02077348, carpo]); Ile Yand��: Houone, [20��03���28������S 164��12���21������E], 7.X.1970, fr., MacKee 22657 (BISH, G, MO, NOU [NOU 048512], P [P 02077238, P 00700823, carpo]); Poum, Anse Pouani, [20��15���42������S 164��05��� 03������E], 0-20 m, 5.I.1971, fr., MacKee 23158 (G, NOU [NOU 048511], P [P 02077347, carpo]); Poum, Anse Pouani, 17.II.1971, fr. & st. fl., MacKee 23348 (G, MO, NOU [NOU 048524], P [P 02077329]); SE de la corne de Koumac, [20��30��� 44������S 164��19���32������E], 250 m, 13.IV.1972, fr., MacKee 25240 (G, P [P 02139399]); Koumac, au N du ruisseau Grand For��t, [20��30���16������S 164��20���45������E], 250 m, 20.IX.1973, fr., MacKee 27477 (NOU [NOU 048514], P [P 02139403, P 02139404], PH); Hienghene, Pindache, [20��42���05������S 165��00���24������E], 100 m, 21.VIII.1976, fr., MacKee 31791 (G, NOU [NOU 048533], P [P 01865133], PH [PH 000 72776]); Thio, Meh, [21��37���44������S 166��15���54������E], 24.VI.1977, fr., MacKee 33347 (NOU [NOU 048510], P [P 01865135], PH); Goro, Nokotar��, [22��19���16������S 167��00���31������E], XI.1978, fr., MacKee 36094 (NOU [NOU 048470], P [P 02095730, carpo], PH [PH 00072768]); Koua Riv., [21��21���51������S 165��42���49������E], 30 m, 4.VII.1979, fr., MacKee 37082 (G, P [P 02077346], PH); Yat��, [22��09���20������S 167��05���53������E], 100 m, 28.VII.1985, fr., MacKee 42691 (NOU [NOU 048536], P [P 02077345], PH); S of Thio: on coastal road, between Nimbo and Camboui Rivers, 21��45���S 166��27���E, 26.IV.1984, fr., McPherson 6514 (MO, PH [PH 0072760]); Port Bouquet, Koum riv., 21��44���21������S 166��22���20������E, 155 m, 24.XI.2011, fr., Munzinger 1040 (MO, NOU); Col de Mo, 22��02���17������S 166��10���48������E, 200 m, 24.XI.2007, fr., Munzinger & al. 4932 (NOU [NOU 050584, NOU 030816]); Tango Plateau, 21��01���18������S 165��00���53������E, 350 m, 16.IV.2008, fr., Munzinger (leg. Butin) 5116 (NOU [NOU 050463]); Creek Pandanus, 21��01��� 29������S 164��46���42������E, fr., Munzinger 5787 (NOU, P [P 00700841, P 00700844]); Ponandou, 20��49���01������S 165��14���04������E, 30 m, 29.IX. 2009, fr., Munzinger et al. 5860 (MO, NOU, P); Creek �� Paul, [20��29���29������S 164��12���29������E], 28 m, 26.XI.2009, fr., Munzinger & al. 6013 (NOU, P [carpo]); Base W du massif de Poum, 20��15���28������S 164��02���40������E, 23.VII.2007, fr., Pillon 758 (NOU [NOU 020033], P [P 02082148, carpo]); Port Bouquet, Ile Toupeti, [21��43���14������S 166��25��� 05������E], 3.XII.1982, fr., Suprin 1919 (G, NOU [NOU048508], P [P02077363], PH); Thio, berge de creek, [21��36���39������S 166��12��� 30������E], 21.II.1972, fr., Veillon 2531 (NOU [NOU 048521, NOU 04 8522, NOU 048523], P [P 02139396]); Thio, [21��36���39������S 166��12��� 30������E], 7.V.1971, y. fr., Veillon 2573 (NOU [NOU 048538, NOU 048 539], P [P 02139397], PH); Extr��mit�� SE de la Pointe N���Go, [22��18��� 40������S 166��41���51������E], XI.1974, fr., Veillon 3129 (MO, NOU [NOU 04 8355, NOU 048356, NOU 048357, NOU 048358, NOU 048359, NOU 048360], P [P 2081996, P 02139456, carpo]); Montagne de Poum, creek au pied du Pic 272, [20��14���55������S 164��01���17������E], 25.III.1982, fr., Veillon 4855 (NOU [NOU 048537, NOU 048443], P [P 02139398, carpo], PH [PH 00072762]); Mont Pani��, [20��33���48������S 164��47���32������E], 300 m, 22.VII.1982, y. fr., Veillon 5249 (NOU [NOU 048478], P [P 02095728], PH); Thio haute Dothio, ca. 200 m, [21��36���39������S 166��12���30������E], 20.X.1988, fr., Veillon 6977 (NOU [NOU 020752], P [P 02077344, carpo])., Published as part of Callmander, Martin W. & Buerki, Sven, 2013, Notes on New Caledonian Pandanaceae: identity and typifications of Pandanus decumbens (Brongn.) Solms and Pandanus reticulatus Vieill., with the new species Pandanus letocartiorum Callm. & Buerki, pp. 51-60 in Candollea 68 (1) on pages 52-54, DOI: 10.15553/c2013v681a4, http://zenodo.org/record/5736361, {"references":["ST. JOHN, H. (1989) Revision of the genus Pandanus Stickman, Part 61. Pandanus of New Caledonia. Harold St John, Honolulu.","IUCN (2001). IUCN Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission, IUCN.","BRONGNIART, A. (1875). Observations sur les Pandanees de la Nouvelle-Caledonie. Ann. Sci. Nat., Bot. ser. 6, 7: 262 - 293.","MCNEILL, J., F. R. BARRIE, H. M. BURDET, V. DEMOULIN, W. GREUTER, D. L. HAWKSWORTH, P. S. HERENDEEN, S. KNAPP, K. MARHOLD, J. PRADO, W. F. PRUD' HOMME VAN REINE, G. F. SMITH, J. H. WIERSEMA & N. J. TURLAND (2012). International Code of Nomenclature for algae, fungi and plants (Melbourne Code) adopted by the Eighteenth International Botanical Congress Melbourne, Australia, July 2011. Regnum Veg. 154.","MARTELLI, U. (1914). Le specie e varieta nuove di \" Pandanus \" menzionate nella enumerazione delle Pandanaceae. Webbia 4: 399 - 435"]}