Nepenthes weda Cheek, 2015

Cheek, M., 2015, Nepenthes (Nepenthaceae) of Halmahera, Indonesia, Blumea 59 (3), pp. 215-225 : 221-223

Nepenthes weda Cheek , sp. nov. — Fig. 2 , 3

Distinguished from N. halmahera in the tepals hairy on the distal part of the adaxial surface (not glabrous); the lower surface of the lid of upper pitchers with a subapical appendage and with the midline free of nectar glands (not without any appendages and with midline with nectar glands). — Type: Bangun et al. 218 sheet 2 of 2 (holotype MO No. 6576095; isotypes BO n.v., MO, photo K), Indonesia, Maluku Islands, N Maluku Province, Central Halmahera, Weda Bay, Bukit Limber, N 00°32'41" E127°58'28", alt. 847 m, male infl., 1 Dec. 2012.

Etymology. Named as a noun in apposition, for Weda Bay in Halmahera, Indonesia, the base of the Weda Bay Nickel Project surveys for which led to the discovery of this species.

Terrestrial climber 2–4 m tall. Stems terete. Short shoots 4–6 mm diam, internodes 1.3–2 cm long, glabrous. Climbing stems 6–9 mm diam, internodes 10.5–14 cm long, axillary buds conical, 2–7 mm long, inserted 8–13 mm above the axil, hairs golden-brown, multicellular, simple, sparse, appressed, 0.5 mm long, mixed with sessile depressed-globose red glands 0.05– 0.1 mm diam. Horizontal stems, subterranean, bearing vertical rosette shoots along their length, 4–6 mm diam, internodes 1.2–4.5 cm long, surface irregularly ridged, corky, glabrous. Leaves petiolate, coriaceous; leaves of nanophyll rosette shoots 3–6, with blades slender linear-oblong, 20–22 by 2.4–6 mm, apex acute, tendril 65–85 mm long, base attenuate, petiole canaliculate 6–10 by 2 mm, base shortly sheathing and clasping the stem by its whole circumference; nerves not conspicuous; both surfaces with scattered glossy pale brown, mainly straight, simple hairs 0.2–0.5 mm long, dense along the adaxial midrib groove, mixed with a few sessile depressed-globose red glands 0.05 mm diam. Leaves of short shoots with blades elliptic-oblong (10.5–)15.2–25.5 by (2.8–)4.1–5.8(–7) cm, apex slightly acuminate, base acute, longitudinal nerves 10 –11 pairs, more or less evenly spread through the lamina, most conspicuous on the lower surface. Pennate nerves numerous, conspicuous above, almost patent, reaching the marginal nerve; margin densely dark brown hairy, hairs simple 0.3–0.6 mm long; upper surface of blade glabrous apart from the densely pubescent midrib, hairs appressed, yellow-brown, 0.5(–0.7) mm long, acute; lower surface with similar hairs covering 20–100 % of midrib, hairs 0.75–1 mm long, red-black or yellow mixed with sessile red glands 0.05 mm diam; inner 1/3 of lamina sparsely hairy, hairs 0.5–1.25 mm long, mixed with sessile red glands, outer 2/3 glabrous or very sparsely hairy. Petiole canaliculate, (1.8–) 3.5–6 cm long, (0.2–) 0.4–0.5 cm wide, the wings of the basal 1–1.5 cm more or less abruptly dilated by c. 5 mm, perfoliate and decurrent for c. 5 mm down the stem, indumentum as in blade in herbarium specimens the dilated wing appearing as a sheath around the stem (artefact of pressing). Leaves of climbing stems with blade very narrowly elliptic, 20–23.5 by 4–4.5 cm, apex rounded-obtuse, base acute-cuneate. Longitudinal and pennate nerves as in the leaves of the short stems; leaf margin indumentum dense, brown, hairs simple, 0.5–1 mm long. Upper surface of blade glossy, glabrescent, initially 10 % covered with simple appressed hairs 0.25–0.6 mm long, mixed with sessile depressed-globose red glands 0.05 mm diam. Lower surface 5–10 % covered in persistent patent pale brown hairs (0.5–)0.75(–1) mm long, and glands as upper surface. Petiole canaliculate, 5–7, 0.4–0.5 cm, base perfoliate-decurrent, clasping the stem for 3/4 its diameter, then decurrent as wings 1–2 mm wide for 1.8–2.7(–4) cm, finally the wings uniting; indumentum as blade. Pitchers of nanophylll rosette shoots placed on ground in leaf litter, matt dark red to red-brown, broadly subcylindric, 4.2–6.5 cm long, 2.5–3.5 cm wide at base, tapering gradually to 1.6–3 cm wide below peristome; fringed wings 1.2–1.5 cm apart, (2–) 7–11 mm broad, fringed elements (1–) 1.8–2.8 mm long, densely clustered, arranged in 3–4 radiating planes, 0.15 mm apart towards peristome, or more evenly spread, 0.8 mm apart in dissection of the wings; indumentum of moderately dense patent simple pale brown hairs, 0.5 mm long, c. 15–20 per mm 2, mixed with red sessile depressed-globose glands 0.03 mm diam at the same density. Mouth orbicular-ovate, slightly broader than long, oblique, slightly concave, column ill-defined, peristome brown-green, rounded in transverse section, 1–1.6 mm wide, ribs more or less conspicuous, 0.4–0.55 mm apart, raised either slightly and inconspicuously or as a short slender wing 0.1 mm high, ribs separated by 12–15 faint striae; outer edge entire, more or less revolute, inner edge extended into conspicuous narrowly triangular, papery teeth 2–2.4 mm long; lid orbicular, 1.2–1.9 cm diam, apex rounded-emar- ginate, base rounded and minutely cordate apex of upper surface with a circular indentation 1–2 mm diam base with 1–3 submarginal tentacles 1–2 mm long; lower surface matt pale yellow, with a corresponding raised, cup-shaped structure, 2 by 1 mm, inset 0.5 mm from apex, the interior densely hairy, otherwise appendages absent, nectar glands absent from midline and margin, 1–3 on each side of the midline, longitudinally elliptic, thickly bordered, domed, glossy yellow, 0.8–1.1 by 0.3–0.8 mm, the central aperture dark glossy brown, elliptic 0.25–0.5 by 0.25 mm, sometimes reduced to a minute pore at slit. Depressed-globose sessile red glands 0.05 mm diam and simple pale brown hairs 0.2 mm long, scattered thinly in the marginal 2–3 mm of the lid, the central portion lacking indumentum completely; marginal 0.6 mm of lid densely hairy, hairs bushy, erect, 0.1 mm long, with 1–3 basal branches; spur entire or trifid, conical, 0.5–1.5 mm long, densely appressed hairy. Lower pitchers (of short stems) brownish maroon, narrowly ovoid-cylindric, 11.3–20.2 cm tall, 4.7–6 cm wide a the basal, ovoid portion, gradually constricting from a point (‘the hip’) 6–7 cm from the base, narrowest below the peristome, 3.6–4.6 cm wide; fringed wings 1.5–2.2 cm apart, 0.9–1.4(–2.8) cm broad, involute, fringed elements 1–2(–3.5) mm long, dense, 0.3 mm apart near the peristome, more widely spaced, c. 1–2 mm apart towards the pitcher base; indumentum of appressed, simple, multicellular, glossy pale brown or red hairs, 0.3–0.6 mm long, covering 10–25 % of the surface; mouth ovate, strongly oblique, straight or very slightly concave, inner surface waxy white with purple blotches, column ill-defined; peristome flattened to rounded green or red, 4–7 mm wide, ribs conspicuous, 0.8–1.1 mm apart, abruptly raised as wings 0.5 mm high, ribs separated by striae; outer edge more or less entire, revolute, inner edge extended into conspicuous, narrowly triangular, papery teeth 2–3 mm long; lid elliptic 3.4–5.9 by 3–4.1 cm, apex rounded, base rounded and minutely cordate; upper surface with indumentum as outer pitcher, sometimes with 3–4 multicellular tentacles 2 mm long, hairy, inserted near the spur (Mahroji 70), purple brown; lower surface with 25–30 thickly bordered, flattened nectar glands (0.6–) 0.75–1.5 mm long, borders glossy, pale yellow, central aperture purple brown 0.2–0.5 mm diam, spur 2–3 mm long, dorsiventrally flat, stout, entire, apex rounded, or (Mahroji 70) fasciculate, with 3 basal, equal branches 1 mm long, densely hairy, hairs 0.25–0.5 mm. Upper pitchers of climbing stems dark red or yellow-green, more or less tinged and mottled red-brown (Phillipson et al. 6417) ovoid-cylindric, 24–25 cm tall, the basal ovoid portion 10 by 6 cm, constricted to c. 4 cm wide above, dilating gradually to c. 5 cm wide below the peristome; wings reduced to two ridges, c. 2 cm apart; indumentum of appressed simple acute hairs as lower pitchers but less dense, and glabrescent in the mature pitcher; mouth ovate, oblique, straight or slightly concave, inner surface waxy white, mottled red, column ill-defined, peristome rounded-flattened in transverse section, 3.5–5.5 mm wide, ribs conspicuous, 1–1.5 mm apart, abruptly raised as wings 0.2–0.3 mm high, ribs separated by striae; outer edge entire, revolute; inner edge extended into concealed, inconspicuous curved teeth 1.5 mm long; lid leathery, elliptic, 6.2 by 4 cm, apex and base rounded, upper surface with indumentum as outer pitcher, lower surface without basal appendage, but with a small subapical appendage 4 – 5 mm long, aligned along the midline, set back 4 mm from the apex, proximal part of appendage bilaterally flattened, convex, semi-circular, 2 by 2 mm, distal part apically flattened, raised 1 mm above the surface, with a central bordered flattened longitudinally elliptic nectar gland c. 1 by 0.5 mm; nectar glands resembling that of the appendage, otherwise scattered on surface except for a midline band c. 5 mm wide at base of lid, widening to 8 mm wide at the subapical appendage; 10–15 nectar glands on each side, orbicular or longitudinally elliptic, orbicular glands 0.5–1.25 mm diam with a central glossy brown centre exposed by an aperture up to 0.5 mm diam in the glossy pale yellow border, elliptic glands 0.75 by 0.5–0.6 mm, larger glands flanking the midline band, smaller glands scattered towards margin, mixed with sparse sessile depressed-globose red glands 0.1 mm diam, marginal 1–2 mm with scattered bushy hairs 0.1 mm diam, surface white, with streaks of red radiating from the basal attachment, nectar glands pale brown in life (Phillipson 6417); spur dorsiventrally flattened, 2–3 by 1 mm, recurved from above half its length, apex rounded, surface minutely densely papillate, papillae translucent, 0.01 mm long. Male inflorescence 44.5–48 by 4.5–5 cm; peduncle 14–18.5 by 0.3–0.4 cm at base; partial-peduncles 65–70, 4-flowered in proximal half, the distal 4/5ths 3-flowered, the distal most 1/5th 2-flowered; bract filiform, re- flexed, 1–4 mm long, inserted c. 2 mm above base; partialpeduncle (3–)5(–7) mm long; partial-rhachis 3–5 mm long; pedicels 8–10(–12) mm long 90 % covered in pale brown appressed simples hairs 0.25 mm long, extending to partial-peduncles, rhachis and peduncle, where covering only c. 20 % of surface. Tepals 4, divided by 9/10, lobes ovate-elliptic, 4–4.2(–4.5) by 3–3.5 mm, apex rounded, upper surface col- oured green, distal half and marginal 0.5 mm with dense appressed simple brown hairs 0.1–0.2 mm long; proximal part glabrous, with minute elliptic-bordered nectar glands 0.05 mm long, c. 0.15 mm apart, at an apex of irregular raised tubercles, black (dried material). Lower surface with dense appressed golden-brown hairs 0.1–0.125 mm long, 90 % cover, androphore terete 2.5–3.5 mm long, 0.5–0.6 mm wide, with scattered simple golden-brown hairs 0.2 mm long; anther head white, subglobose, 1.5–1.6 mm diam, anthers 8 in a single whorl. Female inflorescence, fruit and seed unknown.

Distribution & Ecology — Lower montane forest on ultramafic substrate; 415–1014 m altitude. Only known from Bukit Limber of Weda Bay , Central Halmahera, Indonesia.

Additional specimens. INDONESIA, Maluku Islands, N Maluku Province, Central Halmahera, Weda Bay , km 6 to Bukit Limber, 451 m alt., sterile, 29 Oct. 2012, Mahroji et al. 70 ( BO n.v., MO) ; ibid., Bukit Limber, alt. 1014 m, sterile, 1 Oct. 2012, Phillipson et al. 6430 ( BO n.v., MO 2 sheets); male infl., 25 Sept.2012, Phillipson et al. 6417 ( BO n.v., MO 2 sheets) ; ibid., Bukit Limber, sterile, 20 Apr. 2013, Bidault et al. 1129 ( BO n.v., MO) ; ibid., 4 Dec. 2012, Gushilman 189 ( BO n.v., MO, field image viewed on Tropicos) .

Conservation — Known only from six specimens collected in the course of a botanical survey ( Weda Bay Nickel Project ), in 2012–2013. Described as either common (Phillipson et al. 6417) or uncommon (Phillipson et al. 6430). The evidence available suggests that N. weda is strictly confined to Bukit Limber within the wider project area. This is because while 20 specimens of Nepenthes were collected from seven sites within the c. 23 by 14 km survey area, N. weda only occurred at one of these, Bukit Limber. The nickel and cobalt ore de- posits here are among the deepest and richest known in the zone of the project, and it is planned to remove all vegetation to facilitate open-cast mining in the first phase of the project ( ERM 2010). Given the current evidential basis, this will almost certainly render the species extinct in the wild, unless it can be shown to occur elsewhere, at a safe site. Implementation of the Mitigation Hierarchy in this case appears to be problematic based on current data since N. weda appears to sit on the main orebody (Bukit Limber). Further studies are needed to validate this. Given the higher risk to this species, collection of seed for seedbanking should be considered as a priority.

Therefore, since on current evidence a single location is known with an area of occupancy estimated as 12 km 2 (using the 4 km 2 cells currently favoured by IUCN (2012), and with a similar extent of occurrence, N. weda is here assessed as CR B1+B2a,b(iii), that is Critically Endangered.

The population of N. weda has already almost certainly been reduced by the activities of the Weda Bay Nickel Project , due to their extension of previously existing logging roads in order to access Bukit Limber itself ( ERM 2010).

Population reduction has also been caused by the clearance of vegetation in 2007 for a 12 ha test pit at Bukit Limber, following a vegetation survey ( ERM 2010): “Vegetation was surveyed over 15 ha in the test pit area, slightly larger than the test pit area of 11.3 ha. The survey recorded 31 species of flora, including two protected species of pitcher plants ( Nepenthes sp. and Nepenthes maxima Nees ). The larger surveys of lower montane forest indicate these species are distributed widely within the Contract of Work Area, and elsewhere in Halmahera and Indonesia.... Impacts on the forest are reversible and full recovery is expected in about 20 years.” ( ERM 2010).

It is almost certain that the two Nepenthes taxa referred to in the foregoing passage from ERM (2010) are misidentifications. This is because, subsequent to the initial fieldwork that resulted in ERM (2010), a second survey began in 2012 (and is ongoing) by Weda Bay Nickel , with Missouri Botanical Gardens and Herbarium Bogoriense, that resulted in the specimens identified in this paper as two new species which on the evidence currently available are unique to the Weda Bay Nickel Project area. The high intensity of specimen sampling by the second survey, using Nepenthes as an index, is unprecedented in Halmahera. It can be concluded that the 15 specimen records seen by us from the project footprint, probably represent all the Nepenthes species present within the area surveyed. At Bukit Limber the seven specimens seen belong to N. weda (6 specimens comprising all specimens known globally) and N. halmahera (1 specimen). Contrary to the suggestion in ERM (2010) as quoted above, there is no evidence seen by us to indicate that these species definitely occur elsewhere outside the survey area. While it is credible that N. halmahera which occurs at six out of the seven sites where Nepenthes occur within the survey area, might well occur outside; it is much less likely that N. weda , on current evidence restricted to a single location (Bukit Limber), occurs outside the survey area, although it cannot be entirely ruled out. This may be one of many examples of a point endemic species occurring on ultramafic substrates in SE Asia forests ( Ashton 2014). On current data, another point endemic in the area, which is also assessed as Critically Endangered is Jailoloa halmaherensis (Heatubun) Heatubun & W.J. Baker , a species representing a monotypic palm genus restricted to Gunung Batu, several kilometres to the NE of Bukit Limber, outside of the Weda Bay Nickel Project , in the neighbouring Nickel Mining concession area of Pt Buena Persada (Solway International) ( Heatubun et al. 2014).

Many species of Nepenthes are known in the wild from only a single forested mountain or hill. Examples are N. aristolochioides Jebb & Cheek ( Jebb & Cheek 1997) , N. alzapan Jebb & Cheek ( Cheek & Jebb 2013b) and N. extincta Jebb & Cheek ( Cheek & Jebb 2013h). The last is already thought to be globally extinct due to open-cast nickel ore mining of the ultramafic substrate that forms its habitat ( Cheek & Jebb 2013h).

Reductions in the population of N. weda by the project additional to those caused by the test pit drilling and road clearance have been caused by drilling operations in the exploration phase of the project. These serve to produce cores, which when analysed, give data on the depth of the deposit and the extent of its mineralisation, necessary in order to plan extraction of the ore in an efficient manner. Holes are drilled in ever-intensifying densi- ties. Ultimately, hundreds of holes and their associated drill pads can be expected. Although, commendably, small, lightweight drills have been used in the project, of great benefit in reduc- ing the need for the usual forest clearance, some disturbance of the understorey has still occurred, resulting in damage to N. weda individuals. This can be seen in the photograph of the Wikipedia entry for Halmahera, in the entry for mining, where a small drill rig is shown being moved through a forest, damaging and destroying rosette shoots of what appears to be N. weda (http://en.wikipedia.org/wiki/Halmahera#mediaviewer/ File:Rig_Crew_on _Halmahera _ Island.jpg).

Figures

Fig. 2 Nepenthes weda Cheek. a. Short shoot; with lower pitcher; b. upper pitcher (from climbing stem); c. male inflorescence (lower part); d. perfoliatedecurrent leaf base from climbing stem; e. nanophyll rosette stems from horizontal, rooting main axes, note the reduced leaf-blades and compact pitchers; f. leaf-blade marginal hairs; g. hairs from mid-lateral area, lower pitcher; h. fringed wing of lower pitcher – near peristome; i. spur-lower pitcher; j. lower sur- face of pitcher lid, lower pitcher (submarginal line of dots indicates extent of marginal branched hairs); k. nectar gland detail from lower pitcher; l. peristome, upper pitcher, viewed from above; m. peristome, upper pitcher, viewed from inside pitcher; n. peristome transverse section, inside to right, upper pitcher; o. partial-inflorescence, male flowers; p. details of male flower; q. details of indumentum from lower surface of tepals (a, d, f–k from Bangun et al. 218; b, c, l –q from Phillipson et al. 6417; e from Phillipson et al. 6430). — Scale bars: a–e = 5 cm; f, g = 1 mm; h, i, k–n, p = 2 mm; j, o = 1 cm; q = 0.25 mm. — Drawn by Andrew Brown.

Fig. 3 Nepenthes weda Cheek. Details of pitchers. a. View of inner surface of wing, nanophyll rosette pitcher,showing multiseriate fimbriae and wing nectar gland; b. ibid., lower surface of lid apex,showing ‘U’ shaped appendage; c. ibid., from lower angle of view; d. upper pitcher, lower surface of lid, side view of lid appendage (lid apex to the right); e. upper surface of lower pitcher showing multicellular tentacles (a–c from Phillipson et al. 6430; d from Phillipson et al. 6417; e from Mahroji et al. 70 (drawn from photo on Tropicos). — Scale bars: a–d = 1 mm; e = 1 cm. — Drawn by Martin Cheek.