Brookea tomentosa Bentham. (1876b: 84)

Neo, Louise, Huurne, Suzanne Ter & Adema, Frits, 2019, A taxonomic revision of the genus Brookea (Plantaginaceae), Phytotaxa 403 (3), pp. 171-186 : 180

6. Brookea tomentosa Bentham. (1876b: 84) ( Fig. 5 )

Type: —BORNEO. Sarawak, 1857, Lobb s.n. (holotype K [ K 000979573]).

Shrub or small tree, 1–9 m high, girth 9–30 cm. Bole straight. Bark smooth, lenticellate, light brown, greenish, purplish, dull yellowish or grey; inner bark pale yellow, dull pale yellowish brown, greenish or whitish. Sapwood soft, white or greenish. Old stems may be hollow, inhabited by ants. Indumentum tomentose, rarely apically villous, hairs stellately branched or unbranched. Twigs terete or quadrangular, 4–10 mm diameter. Leaves of a pair unequal, ovate to elliptic, 9–46 by 4–20 cm, index 1.8–2.6, base attenuate, apex acuminate, above green or silvery, thinly to densely tomentose, below densely tomentose, midrib and nerves above flat, nerves 1–14 per side, 9–23 mm apart; petiole 1–6 cm long, grooved, tomentose. Inflorescences 7–27.5 cm long, peduncle 0.5–3.5 cm long. Bracts narrowly ovate, 3–17 mm long, both sides tomentose. Pedicels 2–10 mm long, tomentose. Bracteoles narrowly ovate, 8–10 mm long, both sides tomentose. Calyx tubular, 12–20 mm long, tube 10–14 mm long, teeth of upper lip triangular, 2 by 2 mm, teeth of lower lip triangular, 2 by 2–5 mm, outside tomentose with unbranched and stellately branched hairs, inside thinly sericeous in lower 2/3; in fruit pear-shaped, 13–24 mm long, not torn. Corolla white, sometimes with yellow lines in the throat, pinkish white, greenish or yellowish, 19–38 mm long, tube 15–28 mm long, outside thinly pubescent in upper half, inside glabrous, lobes orbicular, those of upper lip c. 5 by 6 mm, those of lower lip 6–7 by 5–8 mm. Stamens inserted 10–16 mm below the throat, filaments 8–9 mm, resp. 10–12 mm long, glabrous; anthers 1.0–2.1 by 0.5–1.0 mm, glabrous. Ovary ellipsoid, 3–8 by 2–4 mm, glabrous; styles 13–14 mm long, exserted, glabrous. Fruits ovoid, 10–18 by 7–12 mm, (pale) greenish, dark brown or yellow green, glabrous. Seeds 0.4–0.6 by 0.3–0.5 mm, pitted.

Distribution: —Borneo (all states) ( Fig. 2 ).

Habitat: —Primary, secondary, burned or logged forest, riparian forest roadsides, riversides, open areas. Elevation: up to 1000 m. Soil: (yellow) sand, reddish brown silt, sandstone, clay loam.

Phenology: —Flowering and fruiting: throughout the year.

Uses: —In Sarawak, it has been used to clean posteriors of children.

Notes: — Bentham (1876b) described this species very briefly in the notes to B. dasyantha . The branched hairs on the leaf surfaces are more common on the lower surface than on the upper surface. Two specimens (Ambri & Arbainsyah 1723 and Wiriadinata 1186) have an open inflorescence with the flowers well-spaced, and with long pedicels (c. 10 mm long).

Additional specimens examined: —BORNEO. BRUNEI. Belait, Bukit Sawat, Merangking, 60 m, 16 Apr. 1996, Ogata et al. Og-B 130 (BRUN, L), Jalan Merangking-Buau, 4°32’N, 114°35’E, 30 m, 20 Dec. 1994, Suhaili et al. BRUN 16397 (BRUN, L, SAR), Sungai Liang, 4Aug. 1988, Wong WKM 362 (L, SAR, SING), beside Sungai Mendaram, 7 Jun. 1988, Nangkat NN 16 (L, SAR, SING); Labi, Bukit Teraja (4.18 N 114.26 E), start of summit trail, 100 m, 19 Oct. 1991, Simpson & Marsh DS-MM 2040 (L, SING), Labi Hills Forest Reserve, along Bukit Malayan road, 19 Oct. 1995, SATC students et al. BRUN 17172 (BRUN, SING); Lamunin, Ladan Hills Forest Reserve, 5 Aug. 1991, Nangkat NN 216 (L, SAR, SING), 4°38’N 114°46’E, 130 m, 30 Mar. 1991, Sands et al. MS 5702 (BO, L, SAR, SING), 120 m, 9 Jul. 1993, Atkins 446 (BRUN), Sungai Buing, 12 Oct. 2012, Low et al. LYW 389 (SING, SAR); Temburong, en route from Bangar to Sungai Betia, 100 m, 22 Jan. 1964, Hotta 13417 (L); Tutong, Tasik Merimbun, road to Long Mayan, 3 Sep. 1992, Bernstein JHB 84 (BRUN), Merimbun Heritage Park, nature trail around guest house of Merimbun, 4°35’N, 114°40’E, 20 m, 28 Jul. 1999, Suzuki K 12240 (L), Pulau Jelondong, 4°36’N, 114°40’E, 14 Oct. 1989, Forman & Blewett 875 (L, SAR, SING), Simpang Tiga, 31 Mar. 1997, Muzeum staff HA 79 (BRUN), Muara-Tutong highway, at km 40, near the lake, 13 Apr. 1994, Joffre et al. BRUN 15056 (BRUN, L, SAR, SING). INDONESIA. Central Kalimantan: East Kotawaringin, Km 45 from Sangai, c. 1°29’S 112°31’E, 100 m, 27 Sep. 1996, Argent 9633 (BO, L, SING), Permantang, south of Kwala Kwajan, 50 m, 26 Jan. 1954, Alston 13293 (BO, L). East Kalimantan: Berau, km 8 to 35 from Labanan, Berau Forest Management Project (BFMP) Swakelola, Labanan, 2°00’N, 117°00’E, 19 Dec. 1997, Arifin & Ambriansyah Berau 1100 (BO, L), Inhutani area, km 11, trayek A, 50 m, 18 Oct. 1997, Ambriansyah et al. Berau 955 (BO, L), Inhutani I area, near plot 10, 80 m, 7 Oct. 1996, Kessler et al. Berau 141 (L), road to Tanjung Redeb, 2°05’N, 117°17’E, c. 10 km from Tanjung Redeb, 10 m, 14 Feb. 1997, Kessler et al. Berau 258 (L), along road c. 3 km from Labanan (base camp Inhutani I), 2°5’N, 117°17’E, 20 m, 18 Feb. 1997, Kessler et al. Berau 327 (BO, L), road to Tanjung Redeb, c 10 km from Tanjung Redeb, 2°5’N, 117°17’E, 10 m, 14 Feb. 1997, Kessler et al. Berau 261 (BO, L), Labanan, along logging road, Trayek E, 1°58’N, 117°15’E, 200 m, 6 Mar. 2000, Kessler et al. B 1619 (BO); East Kutai, KPC area Bengalon, Kuaria, km 15, 00°50.080’N, 117°34.737’E, 112 m, 9 Apr. 1996, Ambri & Arbainsyah AA 1723 (BO, L), Sebongkok utara, lens 6, 00°50.742’N, 117°32.131’E, 38 m, 17 Apr. 1996, Arifin & Arbainsyah AA 1795 (L), Long Lees, Muara Ancalong, 100 m, 4 Mar. 1978, Wiriadinata 1125 (BO, L), Long Tesak, 100 m, 10 Mar. 1978, Wiriadinata 1186 (BO, L, SING); Kutai Kartanegara, Batu Ampar, Daerah Inhutani I, Bukit Bangkirai, 50 m, 14 Aug. 1993, Marsemi 15 (L), Lua Buluh, 0°14’59”N 116°19’10”E, 31 Aug. 2009, Lee et al. SL 419 (BO, SING), Wanariset, near Samboja, 40 km N of Balikpapan, 6 Dec. 1988, van Balgooy 5711 (BO, L); West Kutai, Long Bagun, 100 m, 24 Jun. 1975, Wiriadinata 571 (BO, L, SING), Soengei Doho, 1896–1897, Jaheri 1593 (BO). North Kalimantan: Bulungan, Sekatak, west of Tarakan, 0 m, 2 Feb. 1979, Murata et al. B 3025 (BO, L), Murata et al. B 3031 (BO); Nunukan, km 40 Perum Perhutani area 1, 4°12’N, 11[6]°46’E, 350 m, 18 May. 2000, Ambriansyah AA 2329 (L), Simenggaris, Dec. 1912, Amdjah 1028 (BO, L); Tarakan, 0 m, 31 Jan. 1979, Murata et al. B 2990 (BO), forest near kg 4, 17 Dec. 1953, Meijer 2574 (BO, L), Tarakan Island, 20 m, 22 Oct. 1953, Kostermans 10738 (BO). South Kalimantan: Barito Kuala, Doeson[?], n.d., Korthals s.n. (L); Baru City, Pulau Laut, 400 m, 13 Dec. 1988, van Balgooy 5747 (L); Tabalong, km 70, PT Aya Yayang Indonesia, 1°39’S, 115°29’E, 300 m, 12 Jul. 2000, Sidiyasa et al. 2086 (BO). West Kalimantan: Bengkayang, Sebalouw [Sebalau], n.d., Teysmann 10888 (BO); Sintang, foot of Gunung Kenepai, 1893, Hallier 1522 (BO, L). MALAYSIA. Sabah: Beaufort, Beaufort Hill, 5°22’N, 115°45’[E], 200 m, 28 Aug. 1983, Beaman et al. 6844 (L), Forest Reserve Singgangau, 20 m, 9 Feb. 1988, Amin SAN 115528 (L), Indian Forest Reserve, Weston, 16 Nov. 1985, Amin SAN 102947 (L, SAR), Jalan Batu 58, 16 Jun. 1988, Amin SAN 115435 (L), Klias, 28 Jul. 1976, Bidin SAN 80742 (L, SAR), Mile 56, Salagon forest, 25 m, 26 Jun. 1992, Amin SAN 126378 (L, SAR), Mile 60 Montinior road, 23 Oct. 1975, Dewol SAN 80245 (L, SAR, SING), Suinggau Forest Reserve, 17 Sep. 1985, Madani & Ismail SAN 111448 (L); Beluran, West of Bukit Luminitong, 11 Mar. 1982, Gibot SAN 94469 (L, SAR, SING), Sungai Sasau, 18 Sep. 1984, Amin & Sigin SAN 67091 (L, SAR, SING); Kalabakan, Bukit Tuku, mile 8 Luasong road, 12 Jul. 1982, Krispinus SAN 95865 (L, SAR), NBT logged area, 8 Apr. 1982, Krispinus SAN 94736 (L, SAR), Ulu Sungai Kalabakan, 17 May. 1984, Sumbing & Matin SAN 103653 (SAR, SING); Keningau, Bornean Sdn Bhd camp, Nabawan, 12 Jun. 1977, Yalip & Ejan SAN 87019 (L, SAR), Kampung Labau, mile [?] 11 Rashna, 26 Aug. 1977, Nordin SAN 85844 (BO, SAR, SING), Rashna area, mile 6, 7 Jul. 1977, Abas SAN 85754 (L, SAR, SING), Sungai Siburan, Sepulot [Sapulut?], 19 Jun. 1986, Leopold et al. SAN 114440 (L), Tambulanan, 22 Oct. 1983, Patrick & Sigin SAN 69094 (L, SAR), 25 Oct. 1983, Amin SAN 68871 (L, SAR, SING), Ulu Sungai Pingas Pingas, 10 Mar. 1988, Mantor SAN 122038 (L); Kinabatangan, Bukit Garam, East of Sungai Mananggol, 20 Feb. 1987, George et al. SAN 117551 (L, SAR); Kota Belud, Mount Kinabalu, Dallas, hill trail near resthouse, 3000 ft, 7 Aug. 1931, Clemens & Clemens 26027 (BO, L); Labuk Sugut, Sungai Kibiribi, 10 Jul. 1983, Gibot SAN 97601 (L, SAR); Lahad Datu, 60 km, Danum valley road, 9 Jul. 1986, Madani SAN 114482 (L); Lamag, Sungai Tangkulap, 28 Jun. 1983, Amin SAN 97422 (BO, SAR, SING), Sungai Ulu Lokan, camp Balat Kanabatangan, 27 Jan. 1983, Patrick & Timbah SAN 95259 (L, SAR); Nabawan, Kampung Tampasison Pandewan/Sapulut, 10 Jan. 1979, Nordin SAN 86021 (SAR), km 55 Jalan Nabawan/Sepulut, Ulu Sungai Nabawan, 21 Feb. 1990, Sumbing SAN 123383 (L), Sungai Pingas-Pingas Maitland area, 26 May. 1986, Asik & Sumbing SAN 115847 (L), Swarikat Undan Sdn Bhd Logging Area, KM18 Jalan Nabawan/ Pandewan, 12 Mar. 1990, Sawan SAN 128065 (L); Papar, Mile 16 Papar-Bongawan road, 380 m, 20 Feb. 1970, Gibot SAN 66724 (L, SAR), Ulu Kimanis, 20 Mar. 1976, Dewol & Talib SAN 80465 (L, SAR); Penampang, at road side km 33 1/2 KK/Tambunan road, 23 Jul. 1984, Amin & Ismail SAN 60461 (SING), 5th mile path from Kampung Babagon to Ulu Terian 5°52’N, 116°11’E app, 2500 ft, 15 Oct. 1969, Cockburn SAN 65494 (L); Pensiangan, Pontomon Forest Reserve, 24 Aug. 1994, Fidilis SAN 139556 (L); Ranau, above Kampung Lawoi, Tampias Telupid, road side, mile 235 Telupid Ranau road, 6 Dec. 1974, Aban & Kodoh SAN 81015 (L, SAR, SING), above Sungai Tampias, 405 m, 4 Feb. 1993, Goh et al. SAN 137103 (L, SAR), southwest of Segindai across Sungai Liwagu, 17 Aug. 1983, Amin SAN 100146 (L, SAR, SING), Ulu Sungai Liwagu, 22 Oct. 1986, Mansus et al. SAN 117154 (L), Ulu Tongod Forest Reserve, Gunung Monkobo Expedition, 5°52’16”N, 116°59’49”E, 196 m, 26 Jul. 2005, Saw et al. SAN 146753 (SAR), side of Sungai Dual, 5 Apr. 1983, Amin et al. SAN 95264 (L, SAR), Ulu Sungai Tungud, 5°52’44”N, 116°58’41”E, 179 m, 26 Jul. 2005, Sang et al. SAN 146212 (SAR); Sandakan, Sandakan and vicinity, Sep. 1920, Ramos 1142 (BO, L), about 0.5 mile, in along trail from mile 79.5, Labuk road, north of road, 21 Aug. 1971, Dewol et al. SAN 74569 (L, SAR), Batu Sapi, 100 ft, 29 Sep. 1963, Sayu SAN 38817 (L, SAR, SING), Batu Sapi road, 19 Aug. 1960, Meijer SAN 21545 (SING), Bode River, 50 ft, 7 Feb. 1935, Keith 4257 (SING), Kebun Cina, 9 Jun. 1990, Meijer SAN 129639 (L), Mile 111 Telupid Ranau road, 300 ft, 7 Aug. 1978, Madani SAN 88853 (SAR, SING), Mile 755 Telupid, 18 Aug. 1979, Gibot SAN 91269 (L, SING), near Camp site, Mile 8 Telupid Ranau road, 8 Mar. 1974, Aban & Saikeh SAN 79335 (L, SAR, SING), Sim Sim Lookout, 500 ft, 21 Feb. 1948, Keith A 1507 (BO, SING), Tangkulap Forest Reserve, landing area, 7 Dec. 1985, Amin SAN 111699 (SAR, SING); Sapulut, Labang, Sepulut [Sapulut?] Forest Reserve, 27 Oct. 1988, Sumbing SAN 125285 (L, SAR, SING); Sipitang, 15 miles from Kampung Mendulong road to Maligan, 4 Sep. 1983, Lee SAN 96859 (L, SAR), Mesapol [Masapol], 30 m, 25 Nov. 1988, Ogata 11670 (L), HS Masapol, 20 m, 18 Aug. 1987, Amin SAN 114723 (L), between Lubuk and Mesapol [Masapol], near Sungai Lakutan, 26 Dec. 1968, Kokawa & Hotta 2694 (L), Kawasan Hutan SFI/ Mendulong, 19 Aug. 1986, Amin SAN 102673 (L), Melalia Forest Reserve, 20 m, 12 Apr. 1988, Amin SAN 115498 (L); Tawau, Benawood logging area, mile 55 Brantian, 17 Jul. 1978, Fedilis & Sumbing SAN 88417 (L), Jalan Hydro, Sepulot, 11 Jun. 1998, Goh & Madani SAN 141162 (L), Logging area, mile 26 Luasong, 21 Feb. 1979, Fedilis & Sumbing SAN 89728 (SAR), Luasong centre old logging area, Ikea planting area, 190 m, 20 Jul. 2001, Postar et al. SAN 144084 (L, SAR, SING), Umas-Umas logging area, 27 Jul. 1977, Krispinus SAN 87299 (L, SAR, SING); Telupid, Sungai Taviu, Ulu Telupid, 18 May. 1984, Amin & Asik SAN 60085 (L, SAR); Tenom, Gunong Lumaku Forest Reserve, 12 Sep. 1991, Maikin et al. SAN 132682 (L), Mendolong, 10 Oct. 1972, Saikeh SAN 72263 (L, SAR, SING), Mandalon Forest Reserve, Sungai Punti, 25 Apr. 1987, Mantor SAN 120348 (L), Tinutudan, Melalap, 15 Jul. 1995, Sumbing SAN 139284 (L), Ulu Tomani, 3 mile from Kampung Kaang, 13 Jun. 1979, Madani SAN 90837 (L); Tongod, Kalabakan, Gunung Rara Forest Reserve, 7 Dec. 1990, Maikin et al. SAN 132161 (L, SAR), west of the base camp, 4°59’30”N, 117°11”E, 100–500 m, 1 Apr. 1997, Sugau et al. JBS 237 (L, SAR), Kampung Saguan, Pinangah, 10 Jul. 1981, Dewol & Patrick SAN 93664 (L), road to Kampung Tamoi, 300 m, 11 Jun. 1983, Dewol SAN 96688 (SAR, SING), Pinangah Forest Reserve, Telupid, 12 Jul. 1993, Sumbing SAN 135651 (L), Sungai Melikop CCL log[ged] area, 350 m, 25 Aug. 1984, Leopold & Dewol SAN 60244 (SAR, SING). Sarawak: n.d., Native collector 2510, 21 Jul. 1954, Wyatt-Smith KEP 80063 (BO, L, SAR, SING); Baram, Geneh Tekuan, upper Geneh River, off Baram, above Long Selatong Ulu, Baram, 750 ft, 19 Jul. 1977, Chin 2777 (L); Belaga, Bakun HEP, Sungai Berangan off Sungai Balui, 510 m, 19 Aug. 1995, Zainudin et al. 5639 (SAR), Ulu Belaga, Batang Belaga, 2°55’N, 113°54’E, 250 m, 31 Oct. 1981, Hansen 924 (L, SAR), Sungai Semawat, 3°00’N, 113°54’E, 250 m, 21 Oct. 1981, Hansen 769 (L, SAR, SING); Bintulu, Trail below summit of Bukit Lumut, Ulu Sungai Lalang, Ulu Sungai Pandan, 720 m, 18 May. 1999, Julaihi et al. 80948 (L), Bukit Mina Wildlife Corridor, south west of Bukit Mina Wildlife Corridor, n.d., Malcolm et al. 90079 (L), 2 Mar. 2007, Malcolm et al. S 96829 (SAR), Sungai Kebalak, Simalajau National Park, 16 May. 1990, Mohtar & Othman 59543 (L, SAR); Kuching, Aug. 1905, Hewitt series 123 (SAR), Oct. 1892, Haviland 2333/1838 (SAR), 5 Jan. 1895, Sarawak Museum s.n. (SING), 19 Apr. 1893, Sarawak Museum s.n. (SING); Lundu, Kubah National Park, road up Mount Serapi, 1°35’37”N, 110°11’7”E, 350 m, 17 Mar. 1995, Beaman et al. 11266 (L), Rambungan, 18 Apr. 2002, Hazlina et al. S 87622 (SAR); Marudi, Lio Matu, upper Baram, 500 ft, 15 Oct. 1920, Moulton SFN 6710 (SING), Logging camp at Tutoh river, ca 10 km N of Long Tarawan, Gunung Mulu National Park, 4°5’N, 114°55’E, 50 m, 29 Mar. 1978, Nielsen 919 (SAR); Miri, Proposed Lambir National Park, 4 Mar. 1966, Morshidi S 24110 (L, SAR, SING), 3 May. 1966, Banyeng S 25054 (L, SAR, SING), southwest Lambir Hills, Lambir formation, 150 ft, 30 Nov. 1962, Au & Ashton S 16779 (BO, L, SAR, SING), Bukit Lambir, 22 May. 1966, Banyeng & Sibat S 24471 (L, SAR, SING), Yong Kow Area, 6½ Mile Bakam road, 200 ft, 6 Mar. 1966, Sibat S 24821 (L, SAR, SING); Nanga Merit, Sungai Lebau, near Rumah Rapek, upper Pelagus, Ulu Kapit, 18 Sep. 1973, Chai et al. S 33218 (L, SAR); Niah, Niah Forest Reserve, 15 Sep. 1982, Ilias & Dami S 43932 (L, SAR); Serian, Sabal Forest Reserve, 250 m, 13 Aug. 1997, Rantai et al. S 76226 (SAR); Siburan, Semengoh Forest Reserve, 24 Jan. 1959, Brunig S 10393 (SAR); Tatau, path to Bukit Kamantan, Ulu Muput Kanan, Anap, 12 Oct. 1963, Chai S 19553 (BO, L, SAR, SING), Lumut Range, Belaga District, Kapit Division, 1 Oct. 1992, Mohtar & Yii S 66214 (L, SAR), Ulu Anap, Sungai Kana, 19 Jun. 1982, Mohtar & Jugah S 44852 (L, SAR).

Figures

FIGURE 2. The distribution of B. tomentosa (white circle ○), B. dasyantha (black circle ●) and B. auriculata (black triangle ▲). Grey shaded areas are 1000 m and above in elevation.

FIGURE 5. Brookea tomentosa Benth. a. Top of twig; b. flower; c. opened corolla; d. opened calyx (corolla removed) (a–d: Ogata et al. Og-B 130). Drawing by Suzanne Ter Huurne.