1. Pimelodendron amboinicum Hassk.

    Pimelodendron amboinicum Hassk., Verslagen Meded. Afd. Natuurk. Kon. Akad. Wetensch. 4 (1855) 140; Pax in Engl., Pflanzenr. IV.147.v (1912) 54; J.J.Sm., Bull. Jard. Bot. Buitenzorg III, 6 (1924) 101; Whitmore, Guide For. Brit. Solomon Ils. (1966) 71; Airy Shaw, Kew Bull. 31 (1976) 398; Kew Bull. Add. Ser. 8 (1980) 196;Kew Bull. 35 (1980) 666; Kew Bull. 36 (1981) 339; Kew Bull. 37 (1982) 34; Djarwaningsih, Blumea 49 (2004) 413, fig. 1a, b, 2a, 3a, b, map 1. — Neotype (designated by Djarwaningsih, 2004): Béguin 2299 (holo BO; iso L, PNH), Moluccas, W Tobelo, Halmahera, see note 1.

    Pimelodendron papuanum Warb., Bot. Jahrb. Syst. 18 (1894) 198; Pax in Engl., Pflanzenr. IV.147.v (1912) 55. — Lectotype (designated by Djarwaningsih, 2004): Hellwig 464 (holo BO; iso B †), New Guinea, Kaiser Wilhelmsland, Finschhafen, bei Butaueng.

    Daphniphyllum conglutinosum Hemsl., Bull. Miscell. Inform. (1895) 137. — Type: Comins 75 (holo K, n.v.), Solomon Islands, San Cristoval.

    ? Pimelodendron naumannianum Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xvii (1924) 200. — Type: Naumann s.n. (holo B, †), Indonesia, Lesser Sunda Islands, Timor, Nordseite der Kupang Bay; see note 2.

Trees, 10–35 m tall, dbh 7–70 cm, bole 5–20 m high. Outer bark blackish brown to red-brown, smooth, not fissured, not peeling; inner bark yellow; sapwood and heartwood white. Leaves: petiole 1–10.5 cm long, apically with transverse ridges adaxially; blade elliptic (to obovate), 6.5–24.5 by 3–12 cm, length/width ratio 2–2.2, coriaceous, base attenuate, usually with a pair of glands abaxially, margin (rarely serrulate to) serrate (to deeply serrate, teeth 2–3 mm long, the tip apiculate, in some collections of Sulawesi), distance between teeth 1.4–3 cm (2–4 cm in some places on Sulawesi), apex acute to cuspidate, upper surface dull when dry; venation indistinct above, prominent beneath, nerves 5–11 pairs. Staminate inflorescences racemose, fascicles up to6 per axil, 1–5.5 cm long, axillary and/or cauliflorous; bracts 3, triangular, apex serrate, central one larger, 1–1.5 by 0.7–1.5 mm, 2 smaller lateral bracteoles c. 0.25 by 0.25 mm. Staminate flowers 1–6.5 by 2–4 mm, yellow; pedicel 0.2–4 by 0.2–1 mm; calyx lobes 2–3.5 by 1–2 mm, margin and apex serrate; stamens 7–29, in 3 whorls, filaments 0.25–1.5 mm long, anthers 0.25–1.25 by 0.25–1 mm. Pistillate inflorescences solitary, 1–1.6 cm long; bracts 3,  triangular, central one large, c. 1 by 1 mm, 2 smaller lateral bracteoles c. 0.5 by 0.5 mm. Pistillate flowers 3–3.5 by c. 1 mm; pedicel 1.5–2 by c.1 mm long; calyx lobes 1.5–2 by 1–1.5 mm, glands at the union of the basal two lobes, margin and apex entire; ovary 1.2–1.8 by c. 1 mm, locules 3 or 4; stigmas lobes pulvinate, flat. Fruits pyriform, 0.8–1.7 cm in diameter, young light green, mature reddish; pedicel 0.6–2 cm long; remnants of stigmas absent. Seeds globose, one per fruit, 1–1.3 by 0.7–1 cm.

    Distribution — Malesia: Sulawesi, Moluccas, Lesser Sunda Islands, New Guinea; Solomon Islands.

(dots in East Malesia)

    Ecology & Habitat — Primary (mixed Dipterocarp) and secondary forest, rain forest pockets in coastal savannah, swamp forest, or edges of rivers; soil: sandy clay, volcanic, lime-stone. Altitude: sea level up to 1000 m. Flowering and fruiting throughout the year.

    Vernacular names — Sulawesi: Amead’a (Tabaur); gito (Tadjiosch); malala’a (Talaud, Salib); sinto (Moena); tambodja (Baree). Moluccas: Ai mulute, mamina (Malay, Ambonese); angalkenga, asare (Otimmer), dee, ereroe (Ambai), espahat (Buru), gawi (Tobelore), goro koeratji (Bacan), kaporaja (Ternate), lalagoe, lemtja, mamare, moi, moreala (Tobela), ngalim, o tigo-tigono, pake (Galela), pekre (Kebar), pokopokor (Malay, Tobelo), posidi (Wajoli), roebi tima (Mangoli), sepahat (Buru). New Guinea: Komkwa (Manikong), kimoko (Tarie), koro (Randawaja), koromi (Wandammen), bepie (Atam), marendom (Biak), diro (Sidei), joem (Mooi), sowgwa (Manikiong). Solomon Islands: Aisubu (Kwara' ae; Whitmore, 1966).

    Notes — 1. The specimen described by Hasskarl was (probably) lost or he might have used living material to describe the plant. No herbarium material could be traced, therefore a neotype was selected.

    2. The differentiating character for P. papuanum is the short pedicel of the staminate flower, less than 1 mm long (as opposed to 0.2–4 mm in typical P. amboinicum). Examination of several characters of many collections, e.g., staminate inflorescences, pedicel, staminate flower, calyx lobes, number of stamens, petiole length, length of leaf, leaf apex, margins, etc. showed a complete overlap between P. amboinicum and P. papuanum, leaving only the pedicel length as sole criterion to separate both species. This character is rather unreliable due to the variation of the pedicel length in the other species of this genus. Therefore, P. papuanum is here regarded as a synonym of P. amboinicum. Perhaps more pistillate material may reveal more differences.

    3. Pimelodendron naumannianum (Timor) is only known from literature, the type is lost. Based on the description it is considered as a (new) synonym of P. amboinicum.

    4. Several collections of P. amboinicum from Sulawesi in three different places: Malili-South Sulawesi (Cel/V-171; Cel/II-461), SE Sulawesi (Prawiroatmodjo & Soewoko 2005), and some unknown places in Sulawesi (Elbert 3308; Kjellberg 1149) have leaves with deeply serrate margins (teeth 2–3 mm long and the tip apiculate), the distance between the teeth is 2–4 cm, as opposed to 1.4–3 cm in all other collections. However, as this is the only difference, no taxonomic separation seems justified.

2. Pimelodendron griffithianum (Müll.Arg.) Benth.

    Pimelodendron griffithianum (Müll.Arg.) Benth. & Hook.f., Gen. Pl. 3 (1880) 331; Hook.f., Fl. Br. India 5 (1888) 468; Pax in Engl., Pflanzenr. IV.147.v (1912) 56; Ridl., Fl. Malay Penin. 3 (1924) 313; Whitmore, Tree Fl. Malaya 2 (1973) 125; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 186; Kew Bull. 32(1977) 80; Djarwaningsih, Blumea 49 (2004) 415, fig. 1e, 2b, 3d, map 1; Welzen, Djarwaningish & Nanakorn in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 507, FIg. 65, Plate XXVII: 3. — Stomatocalyx griffithianus Müll.Arg. in DC., Prodr. 15, 2 (1866) 1142. — Lectotype (designated by Djarwaningsih, 2004, because of the epithet): Griffith s.n. (holo K n.v.), Malacca.

    Pimelodendron borneense Warb., Bot. Jahrb. Syst. 18 (1894) 199; Pax in Engl., Pflanzenr. IV.147.v (1912) 56. — Type: Beccari PB 293 (holo B †; iso BO), Borneo, Sarawak.

    Pimelodendron acuminatum Merr., Philipp. J. Sci., Bot. 11(1916) 74. — Type: Hose 240 (A holo, photo; L iso!), Sarawak, Baram District, Baram.

    Pimelodendron papaveroides J.J.Sm., Bull. Jard. Bot. Buitenzorg III, 6 (1924) 104. — Type: Grashoff 755 (BO holo!), Sumatra, Palembang, Banjoeasin- en Koeboestreken.

Trees 6–25 m tall, dbh10–91 cm, bole 8–12 m high. Outer bark light brown to reddish brown, smooth to rugose, thin, soft; inner bark light brown with pale yellow sap, c. 2 mm thick, with slight fruit smell; sapwood pale yellow, soft. Leaves: petiole 1–9.5 cm long, apically with transverse ridges adaxially; blade elliptic (to obovate), 6.5–18 by 2.5–9 cm, length/width ratio 2–2.6, thin coriaceous, base attenuate, usually with a pair of glands abaxially, margin serrulate, tips apiculate, distance between teeth 1–2.1 cm, apex acuminate to cuspidate, upper surface dull when dry; venation indistinct above, prominent beneath, nerves 6–8 pairs. Flowers yellowish green, calyx greenish brown. Staminate inflorescences racemose, fascicles up to 4 per axil, 1–6 cm long; bracts 3, triangular (bracts of the form P. acuminatum with glands), apex entire, central one large, c. 1 by 1 mm, 2 smaller lateral bracteoles c. 0.25 by 0.25 mm. Staminate flowers 3.8–8.5 by 3–5.8 mm; pedicel 1.5–6 by 1–1.5 mm (see note 4 for exceptional lengths of 5.5–7.5 mm); calyx lobes 2–5.5 by 1.5–3 mm, margin entire; stamens 6–19, in 2 whorls, filaments 0.25–1 mm long, anthers 0.25–1 by 0.25–1 mm. Pistillate inflorescences up to 2 per axil, 2–6 cm long; bracts 1, triangular, c. 1 by 1 mm, apex entire. Pistillate flowers 7.5–12 by 3.8–6 mm; pedicel 3.2–6 by 1–1.5 mm; calyx lobes 2–4 by 1.5–3 mm, with scattered glands at the union of the basal two lobes, margin and apex entire; ovary c. 3 by 2 mm, locules 4–10, stigmas free and straight. Fruits orbicular, 1.2–3 cm in diameter, glabrous, pale green (immature?); pedicel 0.5–3.5 cm long; remnants of stigmas present. Seeds reniform, 3 per fruit, 0.8–1.9 by 0.6–1.2 cm.

    Distribution — Peninsular Thailand (Airy Shaw, 1977), Malay Peninsula, Sumatra, Borneo.

(triangles in West Malesia)

    Ecology & Habitat — Wet forest, secondary forest, pole forest, semi-swamps, along logging roads or on hill slopes; soil dry to wet, sand or clay. Altitude: sea level to 900 m. Flowering: January to April, July to December; fruiting: April to October, December.

    Vernacular names — Sumatra: djeloetoeng pipih, lantjat imbo (Batak, Dairi), lempanai (Palembang), majang ukit (Batak, Toba), njatoh (Malay), panai-panai (Malay), paoeh hoetan (Malay). Borneo: bua mambur, kelampai sitak (Iban), mampoeloet (Dayak), njulir, pulai, tampang (Bekoempai, Sijang), perah ikan (Malay).

    Notes — 1. There is a complete overlap in measurements between P. borneense and P. griffithianum. However, this is mainly based on staminate specimens, more pistillate specimens are required for a well founded opinion about the synonymy of P. borneense. For the time being I agree with Airy Shaw (1975) that P. borneense is a synonym of P. griffithianum.

    2. There are no real differences between P. acuminatum and P. griffithianum. Sometimes P. acuminatum has staminate bracts with glands, but this difference does not merit a special taxonomic status.

    3. Based on different measurements of staminate and pistillate specimens, and the presence of basal glands on the persistent sepals of the fruits no difference between P. papaveroides and P. griffithianum could be found.

    4. Collections from Borneo (Endert 4913, KEP FRI 41342, Kostermans 21385, van Niel 4051) and from Sumatra (Hotta & Okada 1608) have 7–9.5 mm long staminate flowers and pedicels of 5.5–7.5 mm long. These characters are very different from all other specimens, but there is still  anoverlap. Therefore, these specimens are not given a new status.

3. Pimelodendron macrocarpum J.J.Sm.

    Pimelodendron macrocarpum J.J.Sm., Bull. Jard. Bot. Buitenzorg III, 6 (1924) 103; Airy Shaw, Kew Bull. 25 (1971) 551; Whitmore, Tree Fl. Malaya 2 (1973) 126; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 187; Kew Bull. 36 (1981) 339; Djarwaningsih, Blumea 49 (2004) 417, fig. 2c, 3c, map 2. — Lectotype (designated by Djarwaningsih, 2004): Dumas 1509 (holo BO; iso L), Sumatra, Palembang, Rawas.

Trees up to 30 m tall, dbh 20–40 cm. Bark brown, fissured, glabrous. Leaves: petiole 1–8 cm long, apically without transverse ridges adaxially; blade elliptic to obovate, 5.5–20 by 2–7.5 cm, length/width ratio 2.1–2.8, thinly coriaceous, base attenuate, usually without a pair of glands abaxially, margin mostly serrulate (or serrate) with the tips apiculate, distance between teeth 1–2.5 cm, apex cuspidate, upper surface shiny when dry; venation flat above, raised beneath, nerves 6–0 pairs. Staminate inflorescences panicles, 14–22 cm long; bracts 3, triangular, apex entire, central one large, c. 1 by 1 mm, 2 smaller lateral bracteoles c. 0.25 by 0.25 mm. Staminate flowers 3–4 by 2.5–4 mm; pedicel 1–3 by c. 0.5 mm; calyx lobes 3–4 by 2–2.5 mm, margin and apex entire; stamens 15–20, in 2 whorls, filaments c. 2 mm long, anthers 1-2 by c. 0.75 mm. Pistillate inflorescences 7–10 cm long; bracts 1, cup-shaped, c. 3.5 by c. 1.5 mm. Pistillate flowers 17–22 by 6–10 mm; pedicel 6–12 by 3–4 mm; calyx lobes c. 5 by 3 mm, without basal glands, margin and apex entire; ovary c. 9 by 6–9 mm, locules 15–21; stigma lobes indistinct, flat. Fruits orbicular, 5–13 cm in diameter, yellow, with white latex, fragrant; wall c. 2 mm thick; pedicel not seen; remnants of stigmas absent. Seeds reniform, c. 11 per fruit, 2–3 by 1.7–3.4 cm.

    Distribution — Malay Peninsula, Sumatra, Borneo.

    Ecology & Habitat — Forest on low undulating country. Altitude: 5–550 m. Flowering: April, August, September; fruiting: February, July, September.

    Vernacular names — Malay Peninsula: tuba ikan (Malay). Sumatra: balam riedam (Palembang), balam terong (Palembang), dangkoe (Riau, Palembang), gango (Rawas). Borneo: ketong murung (Malay, Dayak).

    Note — Typical for this species are the large fruits with many locules.

4. Pimelodendron zoanthogyne J.J.Sm.

    Pimelodendron zoanthogyne J.J.Sm., Bull. Jard. Bot. Buitenzorg III, 6 (1924) 106; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 187; Djarwaningsih, Blumea 49 (2004) 418, fig. 2d, 3e, map 3. — Lectotype (designated by Djarwaningsih, 2004): Hallier 866 (holo BO; iso L), Borneo.

Trees 10–24 m tall, dbh 5–45 cm, bole c. 12 m high. Outer bark pale brownish, smooth, not fissured, not peeling, c. 5 mm thick; inner bark c. 3 mm thick, with much pale yellow latex; sapwood pale yellowish; wood very hard. Leaves: petiole 3–12.5 cm long, apically with transverse ridges adaxially; blade elliptic, 4–20.5 by 2–10 cm, length/width ratio 2–2.1, thinly coriaceous, base attenuate, usually with a pair of glands abaxially, margin serrate or rarely serrulate with the tips apiculate, distance between teeth 1–3 cm, apex (rarely acuminate to) cuspidate, upper surface dull when dry; venation flat above, raised below, nerves 7–12 pairs. Staminate inflorescences racemose, green when young, fascicles up to 4 per axil, 2–4.5 cm long; bracts 3,  triangular, apex entire, central one large, c. 1 by 1 mm, rarely with a gland at the base, 2 smaller lateral bracteoles c. 0.25 by 0.25 mm. Staminate flowers 4–9.5 by 2–5 mm, yellow; pedicel 2–5 by c. 1 mm; calyx lobes 2–5 by 1.2–3 mm, margin and apex entire; stamens 7–18, in 2 whorls, filaments 0.25-1.5 mm long, anthers 0.5–1 by 0.2–0.5 mm. Pistillate inflorescences solitary, 2.5–3 cm long; bracts 1, triangular, c. 1 by 1 mm. Pistillate flowers c. 6 by 5 mm, yellow; pedicel c. 1.5 by 1 mm; calyx lobes c. 5 by 3 mm, without basal glands, margin serrate; ovary c. 3 by 1 mm, locules 10–15, stigmas connate, recurved. Fruits orbicular, 1.4–6.6 cm in diameter, yellowish, glabrous, wall soft, thin, flesh pale yellow, with abundant milky latex; pedicel 2.5–4.5 cm long; remnants of stigmas present. Seeds ovoid, 2–6 per fruit, 2–2.5 by 1–1.4 cm.

    Distribution — Peninsular Malaysia (Perak), Sumatra (Aceh), Borneo.

(dots)

    Ecology & Habitat — Primary lowland dipterocarp forest, depleted primary forest, disturbed peat swamp forest, on level alluvial flats to small ridges; soil: clay loam, deep sandy clay with stones. Altitude: 30–200 m. Flowering: April to June, August to October; fruiting: July, December.

    Vernacular names — Borneo: Kelampai sitak (Iban).

    Uses — Fruit edible, flesh with sourish odour and with an insipid or sweetish taste

    Note — This species closely resembles P. griffithianum in especially the staminate flowers. Typical are the pistillate flowers with 10-15 connate and recuved stigmas and the ovoid seeds. Also typical are the long petioles, but here there is overlap with other species.

5. Pimelodendron spec. nov.

Djarwaningsih, Blumea 49 (2004) 418, fig. 1c, d, 4, map 3.

Habit unknown. Bark pale brownish; branchlets and twigs terete, pale brownish, smooth, not fissured. Leaves: petiole 4.5–6 cm long, apically without transverse ridges at the adaxial side, pale brownish; blade elliptic, 9.5–12 by 3.5–5 cm, length/width ratio 2.4–2.7, coriaceous, base attenuate, with a pair of glands abaxially, margin serrate, teeth 0.5–1 mm long, curved inward and the apex apiculate, distance between teeth 0.7–1.5 cm, apex cuspidate, upper surface dull when dry; venation flat above, raised beneath. Inflorescences and flowers unknown. Infructescences racemose, 3–4 cm long; bracts and bracteoles not seen. Fruits conical, c. 3 by 2.1–2.5 cm in diameter, 4-locular, glabrous; pedicel 1–1.2 cm long, glabrous; remnants of stigmas present. Seeds not seen.

    Distribution — East Coast of Sumatra.

(triangle in N. Sumatra)

    Notes — 1. The cited collection consists only of leaf and infructescences in one collection. This material resembles P. griffithianum (4–10 locules) very closely, but it is different in its conical fruits and serrate leaf margins. No information is given about the plant or about particular ecological circumstances. The inadequate material and the complete absence of additional information

Excluded species

Pimelodendron dispersum Elmer, Leafl. Philipp. Bot. 1 (1908) 308. — Type: Elmer 7245 (holo PNH †; iso L), Philippines, Leyte, Province of Leyte, Palo = Actephila excelsa (Dalzell) Müll.Arg..