Research Article |
Corresponding author: Nyee Fan Lam ( nflam@yahoo.com ) Academic editor: Yasen Mutafchiev
© 2024 Nyee Fan Lam, Halijah Ibrahim, Yen Yen Sam, Rozainah Mohammad Zakaria, Axel Dalberg Poulsen.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Lam NF, Ibrahim H, Sam YY, Mohammad Zakaria R, Poulsen AD (2024) Three new species of Boesenbergia (Zingiberaceae) from Sabah, Malaysia. PhytoKeys 247: 39-53. https://doi.org/10.3897/phytokeys.247.107961
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Three new species of Boesenbergia, B. bosuangii sp. nov., B. ganaensis sp. nov. and B. gokusingii sp. nov. were discovered in Sabah, Malaysia. Boesenbergia bosuangii is similar to B. stenophylla R.M.Sm. in the narrowly ovate lamina but differs in the shape of the bract and the calyx. Boesenbergia ganaensis is closely allied to B. burttiana R.M.Sm. but differs in the absence of a ligule, the longer petiole, the obtuse leaf base, the acute leaf apex, the bilobed calyx and the anther dehiscing by pores. Finally, B. gokusingii is similar to B. variegata R.M.Sm., by the single leafy shoot but differs in having an unequal, ovate lamina, a cordate leaf base, an acute leaf apex and the anther dehiscing by pores. The three new species are described and illustrated in detail. With the addition of these new species, there are in total 13 species with one variety in Sabah.
Biodiversity, Borneo, endemic, new species, wild gingers
The genus Boesenbergia was initially classified in the tribe Hedychieae in the family Zingiberaceae (
The character, which distinguishes Boesenbergia from all other Zingiberoideae genera, is that the first flowers appear at the top of the inflorescence and flowering progresses towards the base (
Borneo harbours approximately 38 species of Boesenbergia (
Field collections were made between August 2016 and August 2017 at Ranau and Kimanis Districts. The morphology of the new species was analyzed using living plants with reference to herbarium materials (E, K, KEP, KUL, SAN and SING). The procedures of the fieldwork and measurements were conducted based on
Information collected during fieldwork and herbaria was recorded in the Taxon Data Information Sheet (TDIS) form. The form consist of five sections, namely, taxon attributes, geographic range and demographic details on population (
1 | Creeping; leafy shoots normally single-leaved; inflorescence more or less sessile; anther dehiscing by slits | 2 |
– | Erect; leafy shoots with one to many leaves; inflorescence sessile or long pedunculate; anther dehiscing by slits or pores | 7 |
2 | Lamina more or less circular, obtuse or obscurely emarginate at apex | B. orbiculata |
– | Lamina elliptic or narrowly ovate, acute at apex | 3 |
3 | Lamina plain green | 4 |
– | Lamina variegated | 5 |
4 | Floral tube pubescent outside; flower not red at throat; labellum entire | B. flavoalba |
– | Floral tube glabrous outside; flower red at throat labellum bilobed | B. flavorubra |
5 | Petiole 2–3 cm long; lamina 7–12 by 2.5–7 cm, dark green with a band of lighter green up the midrib, variegation sometimes extending to the main lateral veins | B. variegata |
– | Petiole < 0.5 cm long; lamina shorter than 9 cm with width less than 5.5 cm | 6 |
6 | Lamina 4–8 by 1.5–2 cm, mid-green with a broad silver band on either side of the midrib above, surface glabrous | B. kerbyi |
– | Lamina 8.6 by 5.5 cm, upper surface undulating from green to dark green | B. gokusingii |
7 | Fertile shoot single-leaved, rarely bladeless or 2- or 3-leaved | 8 |
– | Fertile shoot with two or more leaves | 12 |
8 | Lamina 50 by 12 cm or larger | B. grandifolia |
– | Lamina much smaller, not exceeding 30 cm long | 9 |
9 | Base of the lamina deeply cordate | B. cordata |
– | Base of the lamina ± attenutate | 10 |
10 | Petiole 17–34 cm long | B. bruneiana |
– | Petiole not exceeding 17 cm | 11 |
11 | Lamina 7–12 cm wide; petiole robust ca. 5 mm thick; lamina with appressed hairs especially around midrib below | B. lambirensis |
– | Lamina less than 7 cm wide; petiole slender, 2 mm or less thick; leaves glabrous | B. ischonosiphon |
12 | Lamina large, much longer than 30 cm | 13 |
– | Lamina shorter than 30 cm, if longer narrower than 7 cm | 17 |
13 | Leaf base thickened with outermost bracts forming a bucket or vase-like structure enclosing inflorescence sometimes together with sheaths of upper leaves; petiole 42–50 cm long | B. jangaruni |
– | Leaf base or sheaths not thickened as above, long-attenuate forming a winged petiole less than 25 cm long | 14 |
14 | Anther dehiscing by subapical pores or slits | 15 |
– | Anther dehiscing by pores | 16 |
15 | Inflorescence densely pubescent; anther ca. 3 mm long, dehiscing by subapical pores | B. hosensis |
– | Inflorescence glabrous, anther ca. 10 mm long, dehiscing by longitudinal slits ca. 6 mm long | B. armeniaca |
16 | Leaf sheath sparsely hairy or glabrous; bracts 5–8 cm; floral tube 8–10 cm; ovary glabrous | B. grandis |
– | Leaf sheath densely hairy; bracts 2–3.5 cm long; floral tube ca. 5.5 cm long; ovary densely hairy in upper half | B. lysichitoides |
17 | Inflorescence long exserted from the leaf sheaths when fully grown, spindle-shaped; flowers red and white | B. pulchella |
– | Inflorescence never long exserted or spindle-shaped; flower colours various | 18 |
18 | Leaves linear; arrangement of blades strongly flabellate | 19 |
– | Leaves elliptic, narrowly ovate or rarely linear to narrowly ovate; arrangement of blades never flabellate | 20 |
19 | Flower plain yellow; anther dehiscing by apical pores; bracts 3.5–6.5 cm | B. flabellata |
– | Flower white, yellow in the centre, pink at the base; anther dehiscing by slits; bracts < 3 cm | B. burttiana |
20 | Leaves variegated | 21 |
– | Leaves plain green | 25 |
21 | Leaves bullate, dark green around main veins and almost silvery on raised area | B. hutchinsonii |
– | Leaves smooth with a silverish or light green central cloud | 22 |
22 | Petiole never exceeding 3 cm, lamina narrowly obovate with attenuate base | B. hirta |
– | Petiole usually much longer than 3 cm, lamina narrowly ovate to elliptic with cuneate base | 23 |
23 | Leaves with a silvery cloud; flowers yellow, labellum orange-spotted | B. ornata |
– | Leaves with yellow central cloud; flowers orange or white with some yellow and reddish purple | 24 |
24 | Leaves 5–12 by 3–4 cm; flower orange, darker at base of labellum; anther dehiscing throughout its entire length | B. aurantiaca |
– | Leaves 18–23 by 4–6 cm flower white with some yellow and reddish purple; anther dehiscing by apical pores, or anther dehiscent only in upper 2/3 | B. belalongensis |
25 | At least a few uppermost leaf sheaths thickened and forming a cup-shaped structure | 26 |
– | Leaf sheath not thickened as above | 27 |
26 | Innermost leaf sheaths enclosing inflorescence much shorter and wider than outer ones; leaves drying darkish brown | B. laevivaginata |
– | All leaves with more or less equal laminae; leaves green or grey-green when dry | B. urceoligena |
27 | Anther dehiscing by slits throughout their length | 28 |
– | Anther dehiscing by pores | 32 |
28 | Petiole < 8 cm | B. imbakensis |
– | Petiole > 10 cm | 29 |
29 | Calyx unilaterally incised (split on one side) | 30 |
– | Calyx tubular | B. sugudensis |
30 | Lamina wider than 4 cm | 31 |
– | Lamina less than 4 cm wide | 33 |
31 | Petiole up to 4 cm long; lamina longer than 10 cm | 32 |
– | Petiole to 2 cm long; lamina 5.2–6.5 by 3.4–3.6 cm | B. truncata |
32 | Lamina 13–16 cm long | B. apiculata |
– | Lamina at least 11.5 by 5.5 cm | B. ganaensis |
33 | Lamina narrowly ovate, more than 12 cm | 34 |
– | Lamina slightly ovate much shorter, up to 12 cm long, if longer, petiole much shorter than 7 cm | 35 |
34 | Lamina 12–20 by 1.5–3 cm; petiole usually to 7–8 cm | B. stenophylla |
– | Lamina 19 by 3 cm; petiole usually at least 10.5 cm | B. bosuangii |
35 | Leaf sheath and ligule hirsute | B. parva |
– | Leaf sheath and ligule almost glabrous | 36 |
36 | Flower yellow-orange | B. oligosperma |
– | Flower white and yellow, occasionally red in throat | B. subulata |
The new species resembles B. stenophylla in having a narrowly ovate lamina, but differs in the shape of the bract (linear elliptic vs. cymbiform) and a tubular calyx (vs. tridentate) (Table
Distinguishing morphological characters of B. bosuangii and B. stenophylla.
Characters | B. bosuangii | B. stenophylla |
---|---|---|
Plant height | to 30 cm | To about 42 cm |
Petiole | 7.5 cm | 10–18 cm |
Leaf apex | Acuminate | Slightly acuminate |
Bracts | 2.7 cm, linear elliptic, pubescent | 4 × 4.5 cm, cymbiform, glabrous |
Calyx | 2-lobed, pubescent | 3-lobed, glabrous |
Labellum | Yellow band from base in the centre spreading towards apex, faint red at base and dark red bands bordering the yellow band, obovate, 1.2 × 0.8 cm | White with yellow centre, flabelliform, 2.5 × 2 cm |
Lateral corolla lobe | White, 1 × 0.25 cm | Pale yellow, 1.5 × 0.4 cm, |
Malaysia. Borneo. Sabah. Cultivated at Kipandi Park, Moyog, 05°54.68'N, 116°06.27'E, 700 m elevation, 8 August 2016, Lam Nyee Fan 356 (holotype BORH!, isotype SAN). Original material collected by Linus Gokusing (BS-23) at Marakau, Gana-gana, Ranau, Sabah, 06°12.24'N, 116°46.03'E, 480–500 m elevation, 3 August 2010.
Terrestrial, evergreen, herb. Rhizome fibrous, subterranean, ca. 1 cm in diameter with 7 cm internodes, white to light brown, base ca. 0.6 cm in diameter, roots white, up to 18 cm long. Leafy shoots 29.5 cm tall, with 1–2 leaves, with 2–3 outer leafless sheaths, 6.0–6.5 × 0.7–1.1 cm, purple brownish, glabrous. Ligule ca. 0.55 cm long, caudate, brown, glabrous. Petiole 6–7.5 cm long, canaliculate, green, reddish at base. Lamina narrowly ovate, 17–17.4 × 2.5–2.75 cm, erect, dark green above, pale green beneath, glabrous, base attenuate, margin entire, apex acuminate, with acumen ca. 2 mm. Inflorescence ca. 2.8 cm, peduncle 1–2 cm, with up to 6 flowers arranged in a one-sided spiral, one flower open at a time. Fertile bracts linear elliptic, ca. 2.7 cm long, white, reddish at apex, pubescent, margin entire, apex attenuate. Bracteole elliptic, ca. 2 × 0.75 cm, translucent, pubescent, margin entire, apex acute. Flower white, born singly from each bract; calyx 1 cm long, tubular, 2-lobed, translucent, pubescent on both surfaces; corolla tube ca. 3.6 cm long, ca. 1.5 mm wide at base, lobes white, glabrous throughout, dorsal lobe ovate-oblong, ca. 1.1 × 0.5 cm, linear elliptic, concave, white, glabrous, apex acute, lateral lobes ovate, ca. 1.0 × 0.25 cm, oblong, glabrous, apex rounded; labellum, obovate, ca. 1.2 cm × 0.8 cm curved-backward, longer than corolla lobes, with yellow band from base in the centre spreading towards lip, faint red at base and dark red bands on both sides, glabrous; lateral staminodes white, reddish at base, linear, ca. 0.6 × 0.25 cm, glabrous; stamen white throughout, ca. 0.45 cm long, filament ca. 4 mm × 1 mm (widest at base), pubescent, anther ca. 0.5 × 0.3 cm, glabrous, anther crest ca. 0.5 × 2 mm, bilobed, pubescent, thecae oblong, ca. 0.3 × 0.1 cm, glabrous, dehiscing by pores; ovary ca. 4 × 1.5 mm, stigma cup-shaped, glabrous; epigynous glands, two, ca. 0.45 cm long, linear, apex truncate, white. Fruit not seen.
Endemic to Borneo, Sabah.
Malaysia. Sabah. Telupid District, Taviu Forest Reserves, c. 200 m elevation, lowland forest, 17 May 2001, Sundaling D, SAN142970 (SING!, SAN!).
The species is named after Dr. Steven Bosuang, owner of Kipandi Park. He is an entomologist doing conservation efforts on insects and plants of Sabah. His collaborations with local and overseas scientist produced many research papers and reports for the conservation of Sabah.
Riverine area in mixed dipterocarp forest, 50–300 m elevation, flowering in August.
Vulnerable VU D1. Boesenbergia bosuangii is endemic to Sabah. This species is found at Ranau and Telupid Districts. Due to the small, restricted populations outside protected area and possible threats from development, landslides and flooding, this species is assessed as VU (Fig.
The new species resembles B. burttiana by having a narrowly ovate lamina and a similar plant height, but differs in having an obtuse leaf base, an acute leaf apex (vs. attenuate leaf base, slightly acuminate leaf apex), absence of ligule, a longer petiole (4 cm vs.1.8 cm) and the anther thecae dehiscing by pores (vs. slit) (Table
Distinguishing morphological characters of B. ganaensis and B. burttiana.
Characters | B. ganaensis | B. burttiana |
---|---|---|
Ligule | Absent | 0.55 cm, acuminate, reddish, glabrous |
Petiole | 4 cm, green, grooved | 1.0–1.8 cm, green, without winged |
Lamina | Upper surface dark green | Upper surface light green |
Leaf size | 11.5 × 5.5 cm | 12–20 × 1.5–3 cm |
Leaf base | Obtuse | Attenuate |
Leaf apex | Acute | Slightly acuminate |
Bracts | 2.4 cm long, white, narrowly ovate, almost translucent | 4 cm long, cymbiform |
Labellum | Yellow band from base in the middle, spreading to almost entire surface near apex with lighter yellow laterally, red bands from 2/3 of the centre yellow band towards side, 1.1 × 0.8 cm | White, with yellow in the centre, pink at the base, 2.5 × 2 cm |
Lateral corolla lobe | Glabrous, white, 1.2 × 0.2 cm, emarginated, white with link pink ad yellowish at base, light pink patches towards lip | Slightly pubescent, pale yellow, 1.5 × 0.8 cm |
Anther | reddish, pubescent | White, glabrous |
Anther dehiscence | Pore | Slit |
Stigma | Truncate | Rounded |
Malaysia. Borneo. Sabah. Cultivated at Kipandi Park, Moyog, 05°54.68'N, 116°06.27'E, 700 m elevation. 8 Aug 2016, Lam Nyee Fan 348 (holotype BORH!, isotype SAN). Original material collected from Ranau, Gana-gana, by Linus Gokusing (BS-15), 05°53.16'N, 116°39.30'E, 700 m elevation, 2 February 2013.
Terrestrial, evergreen, herb. Rhizome fibrous, subterranean, base ca. 0.4 cm in diameter, light brown, roots white, ca. 4 cm long. Leafy shoots ca. 14 cm tall, with 2–4 sheaths, ca. 3.5 × 2.5 cm, glabrous, green, margins entire. Ligule absent. Petiole 2.8–4 cm long, grooved, green. Leafy shoots 1–2. Lamina elliptic, 10–11.5 × 4–5.5 cm, dark green above, green beneath, glabrous, margin entire; base obtuse, apex acute with acumen ca. 1 mm. Inflorescence ca. 2.7 × 0.4 cm, peduncle ca. 0.35 cm with up to 10 flowers arranged in a one-sided spiral, one flower open at a time. Fertile bracts narrowly ovate, ca. 2.4 × 0.5 cm, white, outer and inner surfaces glabrous, almost translucent, margin entire, apex acute. Bracteole elliptic, ca. 1.8 × 0.5 cm, white, outer and inner surfaces glabrous, translucent, margin entire, apex acute. Flower white, born singly from each bract, calyx 0.7 cm long, tubular, white, glabrous, corolla tube white, glabrous, apex rounded, dorsal lobe elliptic, ca. 1.3 × 0.35 cm, concave, lateral lobes elliptic, ca. 1.2 × 0.2 cm, labellum bucket shaped, obovate (when flattened), ca. 1.1 cm × 0.8 cm, yellow band from base in the middle, spreading to almost entire surface towards the apex with lighter yellow laterally with red bands from 2/3 of the centre yellow band towards side, lateral staminodes oblong, ca. 0.7 × 0.2 cm, white, apex acute, glabrous, stamen ca. 1.2 cm long; filament ca. 7 × 1 mm (widest at base), pubescent adaxially and abaxially, anther ca. 3.5 mm long, pubescent; anther crest trilobed, pubescent; thecae oblong, ca. 0.3 × 0.1 cm, white, glabrous, dehiscing by pores, stigma truncate apex, white, glabrous, epigynous glands 0.2–0.4 cm long, linear, apex pointed. Fruit not seen.
Endemic in Borneo, Sabah; known only from the type locality at Kampung [village] Gana-gana, Ranau.
The species epithet refers to the location where the species was collected.
Granite area at 500–600 m elevation.
Vulnerable (VU D2). The species is endemic to Sabah and only found at Ranau, Sabah, Malaysia. There were only four populations found at the site of collection and it has not been found outside the type locality (Fig.
The new species resembles B. variegata R.M.Sm. by having single-leaved shoots and a short petiole, but differs in having unequal ovate lamina (vs. elliptic), a cordate leaf base (vs. rounded or subcordate), an acute leaf apex (vs. subacute) and the anther thecae dehiscing by pores (vs. slits) (Table
Distinguishing morphological characters of B. gokusingii and B. variegata.
Characters | B. gokusingii | B. variegata |
---|---|---|
Ligule length | 1 mm | 2–3 mm |
Lamina | Unequal ovate, upper surface undulating ranging from green to dark green | Elliptic, upper surface dark green with a lighter band along the midrib, sometimes variegated variegation extends to the lateral veins |
Leaf size (cm) | 8.6 × 5.5 cm | 7–12 × 2.5–7 cm |
Leaf base | Cordate | Rounded or subcordate |
Leaf apex | Acute | Subacute |
Bracts | 1.6 cm long, linear to narrowly ovate, glabrous | 0.6–0.8 cm long, ovate, acute, pubescent |
Labellum | With light yellow at base, darker yellow spreading towards lip, obovate, curved-forward, 0.7 × 0.4 cm | Cream with a deep yellow spot in the centre and a deep red spot at the base, elliptic, 1.0 × 0.7 cm |
Lateral corolla lobe | Glabrous, 0.7 cm long | Densely pubescent, 2 cm long |
Anther dehiscence | Pore | Slit |
Malaysia. Borneo. Sabah. Cultivated at Kipandi Park, Moyog, 05°54.68'N, 116°06.27'E, 700 m elevation. 8 Aug 2016, Lam Nyee Fan 361 (holotype BORH!, isotype SAN). Original material collected from Tawau by Linus Gokusing (BS-21b), 05°11.93'N, 117°25.40'E, 500–600 m elevation, 1 Aug 2015.
Terrestrial, evergreen, herb. Rhizome fibrous, subterranean, base ca. 0.4 cm in diameter, roots brown. Leafy shoots ca. 8.5 cm tall, with 1–2 sheaths, ca. 2 × 1.5 cm, glabrous, green, margins entire. Ligule 1 mm long, acute, light brown, glabrous. Petiole 2.8–3.5 cm long, grooved, green. Leafy shoots single. Lamina unequal ovate, 7.5–8.6 × 5–5.5 cm, upper surface undulating ranging from green to dark green, pale green beneath, glabrous, margin entire; base cordate, apex acute with acumen ca. 2 mm. Inflorescence ca. 2.5 × 0.5 cm, peduncle ca. 0.15 cm, with up to 8 flowers arranged in a one-sided spiral, one flower open at a time. Fertile bracts linear narrowly ovate, ca. 1.6 × 0.2 cm, green, pale red at apex, outer and inner surfaces glabrous, almost translucent, margin entire, apex acute. Bracteole linear elliptic, ca. 1.7 × 0.2 cm, white, outer and inner surfaces glabrous, almost translucent, margin entire, apex acute. Flower white, born singly from each bract, calyx 0.4 cm long, tubular, white, glabrous, corolla tube white, glabrous, dorsal lobe narrowly elliptic, ca. 0.7 × 0.2 cm, slightly concave, lateral lobes elliptic, ca. 0.7 × 0.1 cm, labellum obovate, ca. 0.7 cm × 0.4 cm, very light yellow at base, darker yellow spreading towards lip, curved forward, lateral staminodes elliptic, ca. 0.5 × 0.17 cm, white, apex acute, glabrous, stamen ca. 0.75 cm long; filament ca. 4 × 1 mm (widest at base), glabrous adaxially and abaxially, anther ca. 3.5 mm long, glabrous; anther crest rounded, glabrous; thecae oblong, ca. 0.1 × 0.05 cm, white, glabrous, dehiscing by pores, stigma cup-shaped, white, glabrous, epigynous glands 0.1 cm long, linear, apex pointed. Fruit not seen.
Endemic in Borneo, Sabah; known from Tawau and Tambunan.
The species is named after Mr. Linus Gokusing from Kipandi Park. He collected this species from Tawau in 2011. His passion for plants, such as orchids and gingers, has secured information useful to researchers, tourists and botany students.
Mix dipterocarp forest at 500–600 m elevation.
The authors would like to thank the University of Malaya, Universiti Malaysia Sabah, and the Ministry of Higher Education for the funding and research grant for this study. Sincere thanks to the staff of the University of Malaya and Universiti Malaysia Sabah for their assistance in the laboratories and field trips. Our great appreciation to Kipandi Park, Sabah Parks, Sabah Biodiversity Centre for the approvals of access license and collection permits. This study is a part of the first author’s doctoral thesis. Axel Dalberg Poulsen is supported by the players of the People’s Postcode Lottery. The Royal Botanic Garden Edinburgh, is supported by the Scottish Government’s Rural and Environmental Science and Analytical Services Division.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This work was supported by Institut Pengurusan dan Pemantauan Penyelidikan, Universiti Malaya.
Supervision: HI, ADP, YYS. Writing - original draft: NFL. Writing - review and editing: NFL, ADP, RMZ.
Nyee Fan Lam https://orcid.org/0000-0002-6371-4102
Axel Dalberg Poulsen https://orcid.org/0000-0002-7651-6439
The data underpinning the analysis reported in this paper are deposited at GBIF, the Global Biodiversity Information Facility, and are available at https://doi.org/10.15468/gm82es.