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44. Benstonea thurstonii (C. H. Wright) Callm. & Buerki, comb. nova. ≡ Pandanus thurstonii C. H. Wright in Bull. Misc. Inform. Kew 1894: 348. 1894. Typus: FIJI. Viti Levu: V.1894, J. B. Thurston s.n. (holo-: K!; iso-: FI [FI003945]!). Distribution and ecology. – Benstonea thurstonii is endemic to Fiji and has been recorded on the islands of Viti Lev and Koro, and has recently been collected on Vanua Levu (Fig. 6).

22. Benstonea korthalsii (Solms) Callm. & Buerki, comb. nova. Ξ Pandanus korthalsii Solms in Linnaea 42: 12. 1878. Typus: INDONESIA. Kalimatan (Borneo): s.d., Korthals s.n. (holo-: L [L0050555] image seen; iso-: B [B1002 79935, B180002796, carpo.] images seen, L [L0050556, L0050557]images seen). Distribution and ecology. – Benstonea korthalsii occurs in rainforests from Borneo to Sumatra (STONE, 1978).

15. Benstonea elostigma (Martelli) Callm. & Buerki, comb. nova. ≡ Pandanus elostigma Martelli in Boll. Soc. Bot. Ital. 1904: 303. 1904. Typus: MALAYSIA. Perak (Peninsular Malaysia): Larut, IX.1884, King’sCollector (Kunstler) 6559 (lecto-: CAL; isolecto-: SING [SING0059007]!) (designated by ST. JOHN, 1965: 224). Distribution and ecology. – Benstonea elostigma occurs in lowland rainforests at c. 30-700 m in Peninsular Malaysia.

32. Benstonea pectinata (Martelli) Callm. & Buerki, comb. nova. ��� Pandanus pectinatus Martelli in Boll. Soc. Bot. Ital. 1904: 304. 1904. Typus: MALAYSIA. Sarawak (Borneo): Beccari s.n. (holo-: FI [FI003941, carpo.] image seen). Distribution and ecology. ��� Benstonea pectinata occurs in the montane rainforests (900-2500 m) of Borneo and has been collected twice in Sumatra (STONE, 1975a, 1978). Observations. ��� This species is strictly epiphytic and easily distinguished from other epiphytic members of the genus (e.g. B. alticola) by its lobed auricles at the base of the leaves. Benstonea pectinata has often been collected on fallen trees in montane rainforests, but seems to be a strict epiphyte (Fig. 5B)., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 337, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002, {"references":["STONE, B. C. (1975 a). Studies in Malesian Pandanaceae XV. Two new species of Pandanus from Mt. Kinabalu, Sabah (Borneo) and notes on the Pandanaceae of Mt. Kinabalu. Malay. J. Sci. 3: 69 - 74.","STONE, B. C. (1978). Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesiana precursores. Fed. Mus. J. 23: 1 - 74."]}

4. Benstonea andersonii (H. St. John) Callm. & Buerki, comb. nova. ≡ Pandanus andersonii H. St. John in Pacific Sci. 15: 576. 1961. Typus: MALAYSIA. Sarawak (Borneo): Lawas, XI.1960, Anderson s.n. (holo-: SAR!). Distribution and ecology. – Benstonea andersonii is known from freshwater swamp forests of Borneo (Sabah and Sarawak) (STONE, 1993).

21. Benstonea inquilina (B. C. Stone) Callm. & Buerki, comb. nova. Ξ Pandanus inquilinus B. C. Stone in Fed. Mus. J. 23: 60. 1978. Typus: MALAYSIA. Sabah (Borneo): Sandakan Dist., Ulu Dusun, 28.III.1977, Stone & al. 12890 (holo-: KLU!; iso-: K!, PH [PH00018271, PH00018272] image seen, SAN!). Distribution and ecology. – Benstonea inquilina is only known by the type specimen, collected near Sandakan in lowland secondary rainforests.

Key to the genera of Pandanaceae (adapted from STONE,1974; STONE & al., 1998) 1. Pistillate inflorescence (infructescence) a panicle; leaves in 4 spiral rows; flowers with a perianth-like cupule................................................................................. Sararanga 1a. Pistillate inflorescence (infructescence) a cephalium or a raceme or pseudo-umbel of cephalia; leaves in 3 spiral rows; flowers without a perianth-like cupule................. 2 2. Liana; fruit a berry; carpels multi-ovulate (ovules much more numerous than 3); seeds generally free and germinating outside the fruit................................... Freycinetia 2a........................................................................................... Acaulescent shrubs to large trees; fruit a drupe with a hard bony endocarp; carpels uniovulate, rarely 2-3 ovulate; seed bound to endocarp and always germinating within the fruit................................................................ 3 3. Drupes with endocarp not extending between the seed locules, but replaced by the mesocarp extending from the apex to the base; stigmas two, forming a cross at the apex of the pileus; staminodia present at the base of each drupe, each comprising a filament and an anther with four sterile pollen sacs............................................. Martellidendron 3a. Drupes with endocarp extending between the seed locules surrounded by mesocarp; stigma(s) 1 to numerous, of various shapes but never two and forming a cross at the apex of the pileus; staminodia rarely and abnormally present............................................................................. 4 4. Carpels either free or connate into multicarpellate, severalseeded drupes; stigmas always on the adaxial side of the style, variable in shape but usually not sharp and linear (but if so with stigmas always on the abaxial side of the style); staminate flower with several connate stamens, i.e. borne on a stemonophore, rarely free (in P.epipythicus Martelli and P. pentodon Ridl.)...................................... Pandanus 4a. Carpels always free, forming one-seeded drupes; stigmas always positioned on abaxial side of the style, sharp, linear; staminate flower reduced to one stamen, or in triads, free or very slightly joined at the base............. Benstonea, Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on pages 326-327, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002, {"references":["STONE, B. C. (1974). Toward an improved infra-generic classification in Pandanus (Pandanaceae). Bot. Jahrb. Syst. 94: 459 - 540.","STONE, B. C., K. - L. HUYNH & H. H. POPPENDIECK (1998). Pandanaceae. In: KUBITZKI, K. The families and genera of flowering plants 3: 397 - 404. Springer. Berlin.","BUERKI, S., M. W. CALLMANDER, D. S. DEVEY, L. CHAPELL, T. GAL- LAHER, J. MUNZINGER, T. HAEVERMANS & F. F OREST (2012). Straightening out the screw-pines: a first step in understanding phylogenetic relationships within Pandanaceae. Taxon 61: 1010 - 1020."]}

26. Benstonea monticola (F. Muell.) Callm. & Buerki, comb.nova. ��� Pandanus monticola F. Muell., Fragm. 5: 40. 1865. Typus: AUSTRALIA. Queensland: around Rockingham Bay, 1864, Dallachy s.n. (holo-: MEL; iso-: P [P02138 396, P02138397]!). Distribution and ecology. ��� Benstonea monticola is restricted to the lowland rainforests (from sea level to 800 m) in northeastern Australia (Queensland) (WILSON, 2011)., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 336, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002, {"references":["WILSON, A. J. G. (2011). Pandanaceae. In: WILSON, A. J. G. (ed.), Fl. Australia 39: 211 - 235."]}

2. Benstonea affinis (Kurz) Callm. & Buerki, comb.nova. ≡ Pandanus affinis Kurz in J. Bot. 5: 101. 1867. Typus: INDONESIA. BangkaIsl.: Kurz (lecto-: BO) (designated by STONE, 1978: 13). = Pandanus merrillii Warb. in Perkins, Fragm. Fl. Philipp. 1: 50. 1904. Typus: PHILIPPINES. Palawan Isl.: San Antognio Bay, II.1903, Merrill 840 (holo-: B [B100279959] image seen; iso-: FI [FI003946] image seen, NY, US [US00086602] image seen), syn.nov. = Pandanus sarawakensis Martelli in Bull. Soc. Bot. Ital. 1904: 303. 1904. Lectotypus (designated here): MALAYSIA. Sarawak (Borneo): Kuching, VIII.1865, Beccari 350 (FI-B [FI003947: ErbarioBeccari 11902, carpo.]!), syn. nov. Distribution and ecology. – Benstonea affinis is found in peat swamps and gallery forest/river edges from sea level to c. 500 m. It is a widespread species, distributed from Peninsular Malaysia to the Philippines, including Vietnam, Sumatra, Singapore, and Borneo. Observations. –This species is characterized by its 3 to 9 ovoid fruit heads crowded at the apex of an erect peduncle (Fig. 4A). STONE (1972, 1978, 1983a, 1993a), mentionning some differences in dimensions of the fruit, recognized the two species placed in synonymy above, albeit with doubt. He wrote: “ Pandanus merrilli Warb. is so similar to P. affinis Kurz that, on the basis of herbarium material alone, discrimination is virtually impossible” (STONE, 1983a: 208). Pandanus sarawakensis was described by MARTELLI (1904) on the basis of two syntypes: Beccari 350 and 1895. The first of these, misprinted as 550 in MARTELLI (1904: 303), is designated here as the lectotype because it comprises a sheet with a leaf and an infructescence deposited in the Beccari herbarium and a spirit collection of one syncarp in the associated carpological collection. Further taxonomic work is clearly needed for a better understanding of Benstonea affinis with respect to its relatives, especially B. ellipsoidea (Warb.) Callm. & Buerki (see below).

10. Benstonea brunigii (B. C. Stone) Callm. & Buerki, comb. nova. ≡ Pandanus brunigii B. C. Stone in Fed. Mus. J. 23: 32. 1978. Typus: MALAYSIA. Sarawak (Borneo): Miri, Sg. Dalam Forest Reserve, 5.V.1959, E.F .Brunig S17492 (holo-: SAR!). Distribution and ecology. – Benstonea brunigii is endemic to lowland tropical forest understorey in Borneo (Sarawak) (STONE, 1993).

28. Benstonea odoardii (Martelli) Callm. & Buerki, comb.nova. ≡ Pandanus odoardii Martelli in Boll. Soc. Bot. Ital. 1904: 304. 1904. Typus: PAPUA NEW GUINEA. Western Prov.: Fly river, 1877, D’Albertiss.n. (holo-: FI-B [FI003939: Erbario Beccari 1526, 1526A, 1526B] images seen; iso-: FI [FI00 3940] image seen). = Pandanus ihuanus Martelli in J. Arnold Arbor. 10: 141. 1929. Typus: PAPUA NEW GUINEA. Central Prov. : Vailala River, 14.II.1926, Brass 978 (holo-: A [A0002 0041] image seen), syn. nov. = Pandanus rostellatus Merr. & L. M. Perry in J. Arnold Arbor. 20: 182. 1939. Typus: PAPUA NEW GUINEA . CentralProv.: Kubuna, 15.XII.1933, Brass 5675 (holo-: NY [NY00311360] image seen; iso-: A [A0002 0069] image seen), syn. nov. = Pandanus humicola Kaneh. in Bot. Mag. (Tokyo) 55: 305. 1940. Typus: PAPUA NEW GUINEA: rd. side to the Lake Angi, 5.IV.1940, Kanehira & Hatusima 13500 (holo-: FU; iso-: A [A00020040] image seen), syn.nov. = Pandanus eumekes B. C. Stone in Fed. Mus. J. 23: 43. 1978. Typus: INDONESIA. West Papua (New Guinea) : Sorong Dist., Sof Radio Sorong, 7.IV.1954, P. Royen (holo-: L [L0050509] image seen), syn. nov. Distribution and ecology. – Benstonea odoardii, as currently circumscribed, is widely distributed in lowland and montane rainforests of New Guinea (from 100 to 2000 m) (JEBB, 1982). Observations. – The type collection of B . odoardii (D’Albertiss. n.), deposited in the Beccari herbarium (ErbarioBeccari della Malesia) in FI, includes the holotype mounted on three sheets, two with leaves and one with the infructescence. Another possible fragment of the infructescence, considered here as an isotype, is also kept in Martelli’s herbarium. This species is a stemless shrub with long slender leaves (up to 250 cm) and a solitary sub-globose syncarp (c. 6-12 cm in diam.) borne on an erect peduncle (up to 60 cm) (JEBB, 1992). The range of variation of B .odoardii accepted here follows JEBB (1982), who gathered several specimens that cannot be distinguished from other material of this species, even though the styles have a rather variable morphology: either short and acute or more slender to recurved. All this material seem to fall within the currently accepted range of variability of the species (JEBB, 1982). Further collections and fieldwork are required to determine whether this circumscription is appropriate or whether it may include more than a single variable species (JEBB, 1982).

42. Benstonea sylvatica (B. C. Stone) Callm. & Buerki, comb. nova. ��� Pandanus sylvaticus B. C. Stone in Fed. Mus. J. 23: 31. 1978. Typus: MALAYSIA. Sarawak (Borneo): Semengoh, 30. VIII.1977, Stone 13475 (holo-: KLU image seen; iso-: PH [PH00018604, PH00018605] images seen). Distribution and ecology. ��� Benstonea sylvatica is endemic to lowland rainforests of Sarawak (Borneo)., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 339, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002

20. Benstonea humilis (Lour.) Callm. & Buerki, comb. nova. �� Pandanus humilis Lour., Fl. Cochinch.: 603. 1790. Typus: VIETNAM. Tourane [Danang]: Annam, 22.VIII. 1927, J. & M. S . Clemens 4173 (neo-: P [P02085855]!; isoneo-: B, G [G00355114]!, UC) (designated by STONE, 1978: 17). Distribution and ecology. ��� Benstonea humilis grows in rainforests from NE India to Peninsular Malaysia from sea levelto c. 300 m (STONE, 1983b). Observations. ��� Benstonea humilis, as circumscribed here, includes the types ofmany validly published names. Acomplete synonymy can be found in Appendix 1., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 335, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002, {"references":["STONE, B. C. (1978). Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesiana precursores. Fed. Mus. J. 23: 1 - 74.","STONE, B. C. (1983 b). Pandanaceae. In: AUBREVILLE, A. & J. - F. LEROY (ed.), Fl. Cambodge Laos Vietnam 20: 1 - 48. Museum national d'Histoire naturelle, Paris."]}

13. Benstonea copelandii (Merr.) Callm. & Buerki, comb. nova. ≡ Pandanus copelandii Merr. in Publ. Bur. Sci. Gov. Lab. 17: 7. 1904. Typus: PHILIPPINES. Negros: Gimagong River, 5.I.1904, Copeland 140 (PNG destroyed; iso-: FI not located). Neotypus (designated here): PHILIPPINES. Sorsogon Prov. (Luzon): Mt. Bulusan, Irosin, IV.1916, fr., Elmer 15267 (G [G00353830]!; isoneo-: FI [FI003948] image seen, K, L, MO!, P [P01754716]!, U, UC [UC273333] image seen, US [US01115658] image seen). = Pandanus copelandii var. panchoi B. C. Stone in Fed. Mus. J. 23: 25. 1978. Typus: PHILIPPINES. Sorsogon Prov. (Luzon): Gubat, 25.X.1960, fr., Pancho & Bardenas 4820 (holo-: CAHP), syn. nov. Distribution and ecology. – Benstonea copelandii is endemic to the Philippines (ranging from Luzon to Mindanao). Observations. – The holotype, deposited at the herbarium in Manila (PNG), was destroyed during World War II, but a fragment has been located in FI. The original collection was rather incomplete, comprising only a few drupes, which were illustrated in MARTELLI (1914: Tab. 31, Fig. 5- 7) and ST. JOHN (1969: 357, Fig. 295). We have not been able to locate the duplicate material at FI despite an extensive search. Aneotype (Elmer 15267) is therefore designated here. This collection is complete and well represented in several herbaria. Pandanus copelandii var. panchoi is considered as a synonym. It was described on the basis of a collection bearing only a single syncarp (vs 3-8 crowded, spicate syncarp in the remaining collections of Benstonea copelandii), but is considered to fall within the morphological variation of the species, even though it is rather atypical and is documented only by a single collection.

16. Benstonea foetida (Roxb.) Callm. & Buerki, comb.nova. ≡ Pandanus foetidus Roxb., Fl. Ind. ed. 1832, 3: 742. 1832. ≡ Pandanus wallichianus Martelli in Webbia 4: 36. 1913 [nomen nudum]. Neotypus (designated here): BURMA: Amherst, 1827, Cat. Wallich 8588 (K [K000781275]!; isoneo-: K [K000781 276]!, M [M0175808, M0175809, M0175813] images seen). Distribution and ecology. – Benstonea foetida is found in lowland and swamp forests around the Bay of Bengal in Bangladesh, Burma and India (STONE, 1978). Observations. – ROXBURG described this species in 1832 in his Flora Indica. He wrote that it grew wild in Bengal and was cultivated at the Royal Botanic Gardens in Calcutta. A careful search of various herbaria known to hold collections made by Roxburg failed to reveal any authentic material of Benstonea foetida. In WALLICH’ s catalogue (1831), published the year before the protologue, two collections can be attributed to Pandanus foetidus: Wallich 8588 & 8591. The latter of these represents a staminate inflorescence (E [E00567756] image seen; FI image seen; Kimage seen; M [M0175810] image seen) from Sillet [Sylhet] in Bangladesh. The other collection, Wallich 8588, consists of leaves and separate drupes from Amherst in Burma (WALLICH, 2005). This collection, held at Kwith some possible duplicates in M, was cited under the nomen nudum, P.wallichianus (MARTELLI, 1913). The Kspecimen is here designated as the neotype.

19. Benstonea herbacea (Martelli) Callm. & Buerki, comb. nova. �� Pandanus herbaceus Martelli in Boll. Soc. Bot. Ital. 1904: 303. 1904. Lectotypus (designated here): MALAYSIA. Perak (Peninsular Malaysia): Trang, III.1881, Kunstler 1397 (FI [FI003936]image seen). Distribution and ecology. ��� Benstonea herbacea grows in lowland and mountain rainforests (600-1300 m) of northern Peninsular Malaysia and the southern part of Thailand (STONE, 1978). Observations. ��� The collection locality for the type of Pandanus herbaceus Martelli was cited as ���Irang, Penins. Malese (H. Calc.)��� (MARTELLI, 1904: 303). STONE (1978: 37) later indicated ��� coll. ignot. ���, but a fragment in Martelli���s herbarium (FI) clearly comes from the type locality cited by Martelli and was collected by H. H. Kunstler (Kunstler 1397),and should be considered as typema terial.Because the holotype inCAL was not located by B. C. Stone and appears to have been lost, the collection in FIis here designated as the lectotype., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 335, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002, {"references":["STONE, B. C. (1978). Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesiana precursores. Fed. Mus. J. 23: 1 - 74.","MARTELLI, U. (1904). Pandani asiatici nuovi. Bull. Soc. Bot. Ital. 1904: 298 - 304."]}

30. Benstonea pachyphylla (Merr.) Callm. & Buerki, comb. nova. ��� Pandanus pachyphyllus Merr. in J. Straits Branch Roy. Asiat. Soc. 85: 154. 1922. Lectotypus (designated here): MALAYSIA. Sabah (Borneo): Sandakan and vincity, IX-X.1920, fr., Ramos 1541 (US [US00086615] image seen; isolecto-: K [K00078 1368, K000781386]!, P [P02138353]!). Distribution and ecology. ��� Benstonea pachyphylla is endemic to the lowland rainforests of Borneo (Kalimatan, Sabah and Sarawak). Observations. ��� This species is very characteristic, with its acaulescent habit and syncarp borne in the middle of a large clump of leaves growing directly from the ground (Fig. 4D). Its leaves are similar to those of the poorly known B.celebica, the latter only known by its leaves and a picture of the fruit (see above). These may represent a single species that occurs in both Sulawesi and Borneo., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 337, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002

25. Benstonea lepatophila (B. C. Stone) Callm. & Buerki, comb. nova. ��� Pandanus lepatophilus B. C. Stone in Bot. J. Linn. Soc. 85: 31. 1982. Typus: MALAYSIA. Sarawak (Borneo): Gunung Mulu NP, 14.IV.1978, Argent & Jermy 973 (holo-: E [E00318031] image seen; iso-: KLU!, SAR!). Distribution and ecology. ��� Benstonea lepatophila is endemic to the limestone hills of Gunung Mulu National Park in Borneo (Sarawak) (STONE, 1993)., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 336, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002, {"references":["STONE, B. C. (1993). Studies in the Malesian Pandanaceae 21. The genus Pandanus in Borneo. Sandakania 2: 35 - 84."]}

23. Benstonea kurzii (Merr.) Callm.& Buerki, comb.nova. �� Pandanus kurzii Merr., Interpr. Herb. Amboin.: 107. 1917. Typus: INDONESIA. Java: Mt. Salak, 17.V.1863, Coll. ignot. (lecto-: BO) (designated by STONE, 1972: 29). Distribution and ecology.��� Benstonea kurzii is confined to the rainforests of Java (Indonesia)(STONE, 1978)., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 335, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002, {"references":["STONE, B. C. (1972). Studies in the Malesian Pandanaceae X. Four new Malesian species of Pandanus and observations on P. sarawakensis. Fed. Mus. J. 14: 127 - 136.","STONE, B. C. (1978). Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesiana precursores. Fed. Mus. J. 23: 1 - 74."]}

17. Benstonea gibbsiana (Martelli) Callm. & Buerki, comb. nova. ��� Pandanus gibbsianus Martelli in J. Linn. Soc., Bot. 42: 170. 1914. Typus: MALAYSIA. Sabah (Borneo): between Tambunan and Korikut, II.1930, S. L . Gibbs 3030 (holo-: BM [BM000958510]!). Distribution and ecology. ��� Benstonea gibbsiana is known only from lowland rainforests (> 800 m) in Sabah (Borneo) (Fig. 4B)., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 333, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002

34. Benstonea pilaris (Ridl.) Callm. & Buerki, comb.nova. ��� Pandanus pilaris Ridl. in J. Straits Branch Roy. Asiat. Soc. 86: 311. 1922. Lectotypus (designated here): MALAYSIA. Negeri Sembilan (Peninsular Malaysia): Bukit Tangga, 10.XII.1920, Ridley s.n. (K [K00781286]!). Epitypus (designated here): MALAYSIA. Selangor (Peninsular Malaysia): Pantai Valley, 13.IX.1967, Mahmud & Sharif s.n. (KLU image seen). Distribution and ecology. ��� Benstonea pilaris is endemic to the rainforests of Peninsular Malaysia (STONE, 1969). Observations. ��� The type collection (Ridley s.n.) has not been found in SING, but a steril duplicate has recently been rediscovered by HB at Kew, which is here designated as the lectotype. In 1978, STONE designated a specimen at KLU (Mahmud & Sharif s.n.) as the neotype, but the rediscovery of the type revokes this nomenclatural act. As Mahmud & Sharif s.n. is a fertile collection including a complete syncarp, and clearly belongs to B.pilaris, we designate it as the epitype., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 338, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002, {"references":["STONE, B. C. (1969). Studies in the Malesian Pandanaceae, VI. The identity of \" Pandanus pilaris \" Ridley. Webbia 23: 609 - 613.","STONE, B. C. (1978). Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesiana precursores. Fed. Mus. J. 23: 1 - 74."]}

Callmander, Martin W., Lowry Ii, Porter P., Forest, Félix, Devey, Dion S., Beentje, Henk, Buerki, Sven (2012): Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species. Candollea 67 (2): 323-345, DOI: 10.15553/c2012v672a12

50. Benstonea vriensii (Martelli) Callm. & Buerki, comb. nova. ��� Pandanus vriensii Martelli in Webbia 4: 433. 1914. Typus: INDONESIA. Sumatra: Battakbergen, I.1905, De Vriens s.n. (holo-: FI [FI003944: 3 sheets] images seen). Distribution and ecology. ��� Benstonea vriensii is confined to the lowland rainforests of Sumatra (Indonesia) (STONE, 1978)., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 341, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002, {"references":["STONE, B. C. (1978). Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesiana precursores. Fed. Mus. J. 23: 1 - 74."]}

49. Benstonea unguiculata (Ridl.) Callm. & Buerki, comb. nova. ��� Pandanus unguiculatus Ridl., Mat. Fl. Malay. Penins. 2: 229. 1907. Typus: MALAYSIA. Selangor (Peninsular Malaysia): Gunung Kutu, V.1896, Ridley 7659 (holo-: SING [SING00 59105]!). Distribution and ecology. ��� Benstonea unguiculata is endemic to lowland rainforests of Peninsular Malaysia from sea levelto 1500 m (STONE, 1978)., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 341, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002, {"references":["STONE, B. C. (1978). Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesiana precursores. Fed. Mus. J. 23: 1 - 74."]}

27. Benstonea nana (Martelli) Callm. & Buerki, comb.nova. ��� Pandanus nanus Martelli in Webbia 1: 370. 1905. Typus: MALAYSIA. Perak (Peninsular Malaysia): Larut, IV.1883, Kunstler 4080 (holo-: FI [FI003936: 3 sheets] images seen; iso-: SING [SING0059057]!). Distribution and ecology. ��� Benstonea nana is endemic to the lowland rainforests of Peninsular Malaysia (Perak and Pahang) and southern Thailand occurring at an altitude of 90- 300 m (STONE, 1978)., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 336, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002, {"references":["STONE, B. C. (1978). Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesiana precursores. Fed. Mus. J. 23: 1 - 74."]}

38. Benstonea rupestris (B. C. Stone) Callm. & Buerki, comb. nova. ��� Pandanus rupestris B. C. Stone in Fed. Mus. J. 14: 129. 1972. Typus: MALAYSIA. Sarawak (Borneo): Bukit Pantu, Melinau, Kapit, 9.VIII.1967, llias Paie 25728 (holo-: SAR!). = Pandanus oresbios B. C. Stone in Fed. Mus. J. 23: 58. 1978. Typus: INDONESIA. Kalimatan (Borneo): Central E, near Mt. Kemul, 12.X.1925, Endert 3896 (holo-: L [L0050606] image seen; iso-: A [A00019993] image seen), syn. nov. = Pandanus scandens B. C. Stone in Fed. Mus. J. 23: 59. 1978. Typus: MALAYSIA. Sarawak (Borneo): Bintulu, Merurong Plateau, 16.IV.1960, Brunig 29924 (holo-: SAR!), syn. nov. Distribution and ecology. ��� Benstonea rupestris is endemic to the lowland rainforests of Sabah and Sarawak (Borneo). Observations. ��� STONE (1978) described Pandanus scandens based on a more mature infructescence than that of P. rupestris, which was described six years earlier. He distinguished these two taxa based on the number of syncarps (5 in P.scandens vs 7 to 9 in P. rupestris) and other minor morphological characters such as the leaf length (c. 3 �� 105 cm vs 3-4 190 cm) and color (glaucous vs not glaucous beneath). Recent observations in the field by MWC and SB have, however, confirmed that populations corresponding to Benstonea rupestris are characterized by along pendent infructescence with up to 9 oblong syncarps (sometimes 5 to 8) and that leaf length and width fall within the normal variation of B.rupestris as currently circumscribed (Fig. 5C). Pandanus oresbios, also described in 1978, is here considered as a synonym of Benstonea rupestris. It was described based on an immature infructescence, and despite its apparent narrower leaves (2 cm vs 3-4 cm in B. rupestris) and smaller syncarps (c. 3.5 cm in length vs 8 cm), it appears to represent the same entity as B.rupestris. Further field observations are needed to understand the relationship between B.rupestris and B.calcinacta, another species that may also be part of this group but is provisionally accepted here (see above)., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on page 338, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002, {"references":["STONE, B. C. (1978). Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesiana precursores. Fed. Mus. J. 23: 1 - 74."]}

Benstonea Callm. & Buerki, nom. & stat. nov. ��� Pandanus sect. Acrostigma Kurz in J. Bot. 5: 100. 1867. ��� Pandanus subg. Acrostigma(Kurz) B. C. Stone in Bot. Jahrb. Syst. 94: 521. 1974 [non Acrostigma O. F. Cook & Doyle (Arecaceae) in Contr. U.S. Natl. Herb. 16: 228. 1913]. Type: Benstonea affinis (Kurz) Callm. & Buerki (��� Pandanus affinis Kurz) (lectotype designated by ST. JOHN 1960: 226). Acaulescent or short-stemmed shrubs, often epiphytic, rarely tall trees. Leaves ligulate to linear-attenuate; leaf apex adaxially spinulose along the two main pleats (spines rarely absent). Inflorescences terminal or lateral on short side-branches, pistillate cephalia solitary, sometimes spicately disposed. Cephalia always of simple drupes, drupes never connate into phalanges; pileus usually distinct and calyptrate, grading upward into a hard, spiniform style; stigma linear, always positioned on abaxial side of the style. Endocarp usually with seed-chamber roofed by a thin cartilaginous partition above which is located a small more or less distinct supra-seminal chamber distinct from the rest of the apical mesocarp. Staminate inflorescence normally spicate. Staminate flowers sessile, composed of free stamens with anthers much longer than the short or nearly obsolete filaments, apiculate, sometimes stamens arranged in pauci-staminate dyads, phalanges or triads. Etymology.��� This genus is named in honour of Benjamin Clemens Stone (1933-1994) to whom we dedicate this article. Ben contributed immensely to the taxonomy of Pandanaceae and was the leading authority on the family for several decades; his work will remain fundamental to the study of the group for many years to come. Observations. ��� As suggested above, our current understanding of relationships indicates that the entity recognized by STONE (1974) as Pandanus subg. Acrostigma is probably not monophyletic, and that the group treated here as Benstonea should be limited to include only the members historically assigned to Pandanus sect. Acrostigma (excluding P.microglottis), along with two clearly related species originally placed in sect. Fusiforma (P.pachyphyllus and P. biplicatus). The other species assigned to Pandanus subg. Acrostigma by STONE (1974), viz. the members of sects. Epiphytica and Pseudoacrostigma, along with the remaining species of sect. Fusiforma, are morphologically distinct and currently remain in Pandanus (as redefined here), although their taxonomic affinities within the family remain to be elucidated. Benstonea is the third largest genus of Pandanaceae, with 50 currently accepted species (see below). It ranges from India to Australia with its centre of diversity in Borneo (21 spp. [17 of which are endemic]) and Peninsular Malaysia (14 spp. [6 endemic]), where two third of the species are found (Fig. 3). Most members of the genus are acaulescent or shortstemmed shrubs, or facultative or true epiphytes, rarely tall trees [e.g. B. atrocarpus (Griff.) Callm. & Buerki]. Species occur in a variety of habitats including lowland rainforest, swamps, mangrove fringes and montane forests on various soil types (granitic, sandstone and limestone) (STONE, 1978). Benstonea can be easily recognized in the field by its: 1) staminate flowers bearing single (sometimes 2 or 3 connate) stamens with a short filament and a narrow anther; and 2) pistillate drupes always monocarpellate, with a sharp spiniform style and an abaxial stigmatic groove (Figs. 4-6). When fertile, the only taxa with which Benstonea can possibly be confused are some species of Pandanus sect. Solmsia B. C. Stone from subg. Rykia (de Vriese) B. C. Stone, which also bear simple spiniform styles, but which have a stigmatic groove that is always adaxial on the style. The presence of teeth along the two ventral pleats of the leaf apex can also help to recognize Benstonea when sterile, although some Pandanus species found in regions where Benstonea is present (i.e. Pandanus echinodermos Holttum & H. St. John and P.pentodon Ridl., both occurring in Peninsular Malaysia) also bear ventral apical teeth (STONE, 1968a)., Published as part of Callmander, Martin W., Lowry Ii, Porter P., Forest, F��lix, Devey, Dion S., Beentje, Henk & Buerki, Sven, 2012, Benstonea Callm. & Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species, pp. 323-345 in Candollea 67 (2) on pages 328-329, DOI: 10.15553/c2012v672a12, http://zenodo.org/record/5790002, {"references":["ST. JOHN, H. (1960). Revision of the genus Pandanus Stickman, Part 1. Keys to the sections. Pacific Sci. 14: 224 - 241.","STONE, B. C. (1974). Toward an improved infra-generic classification in Pandanus (Pandanaceae). Bot. Jahrb. Syst. 94: 459 - 540.","STONE, B. C. (1978). Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesiana precursores. Fed. Mus. J. 23: 1 - 74.","STONE, B. C. (1968 a). Pandanus Stickm. in the Malayan Peninsula, Singapore, and lower Thailand. Part IV. Malay. Nat. J. 21: 125 - 141."]}