Nepenthes fusca auct Kurata (1976: 48) Phillipps & Lamb (1988: 24) Phillipps & Lamb (1996: 87) Jebb & Cheek (1997: 41) Clarke (1997: 87) Cheek & Jebb (2001: 64) Phillipps et al. (2008: 121) McPherson (2009: 346) Nepenthes maxima auct Kondo & Kondo (1983: 110) Kondo & Kondo (2006: 121) Revisions in Nepenthes following explorations of the Kemul Massif and the surrounding region in north-central Kalimantan, Borneo Robinson, Alastair S. Golos, Michal R. Barer, Marc Sano, Yosuke Forgie, Jennifer J. Garrido, Diego Gorman, Chandler N. Luick, Adi O. Mcintosh, Nick W. R. Mcpherson, Stewart R. Palena, Gregory J. Pančo, Ivan Quinn, Brian D. Shea, Jeff Phytotaxa 2019 2019-02-14 392 2 97 126 73X9X (J. H. Adam & Wilcock) J. H. Adam & Hafiza J. H. Adam & Hafiza J. H. Adam & Wilcock [165,897,1274,1301] Magnoliopsida Nepenthaceae Nepenthes Plantae Caryophyllales 17 114 Tracheophyta species zakriana  Golos ≡  Nepenthes curtisiisubsp. zakrianaJ.H. Adam & Wilcock (1998: 151, fig. XXIIa–b) ( Figs. 9−10)   Type:—Borneo, Sabah, Mt. Kinabalu, Mamut, 1100 m, 21 January 1988, Adam, Adam & Aliosman 2431( holotypeUKMB, iso-ABD). − “  Nepenthes stenoperculum” ined .[annotated on the specimen (Clemens & Clemens) 30980/31092, deposited at BO].    − Nepenthes fusca auct. nonDanser  Kurata (1976: 48), partim[pl. 13]; [  Phillipps & Lamb (1988: 24), partim[3 figs.];  Phillipps & Lamb (1996: 87), partim[fig. 48];  Jebb & Cheek (1997: 41), partim;  Clarke (1997: 87), partim[fig. 57];  Cheek & Jebb (2001: 64), partim;  Phillipps et al.(2008: 121), partim[figs. 149–150, 152–153];  McPherson (2009: 346), partim[figs. 183–185]].   − Nepenthes maxima auct. nonReinw. exNees [  Kondo & Kondo (1983: 110), partim[1 fig.];  Kondo & Kondo (2006: 121), partim[1 fig.]].   Description 2:—Epiphytic or terrestrial climbing or scrambling shrub, to 3 mtall. Stemsof rosettes terete, 0.4–0.6 cmin diameter, internodes 0.5–1.5 cmlong; vining stems ca. 0.5 cmin diameter, internodes 6–15 cmlong. Leaves of rosettescoriaceous, petiolate, elliptic to obovate, 10–12 cmlong, 2.5–4.5 cmwide, apex acute to obtuse, base gradually attenuate into petiole, petiole 1.5–4 cmlong, base amplexicaul and sheathing stem by 1/2 to 4/5 its circumference, not decurrent to slightly decurrent for less than 8 mm. Longitudinal veins 2–3 on either side of midrib in outer third of lamina, running parallel to the laminar margin, originating at base of leaf, inconspicuous. Pinnate veins numerous, irregularly reticulate, indistinct. Tendrils ±2 times the length of the lamina, generally 15–20 cmlong. Leaves of climbing stemsscattered, coriaceous, petiolate, lamina narrowly elliptic to elliptic, rarely obovate, 9–16 cmlong, 3.5–5 cmwide, apex acute, base gradually attenuate, petiole 3.5–6 cmlong, base amplexicaul, sheathing stem by 1/2 to 4/5 its circumference, not decurrent to slightly decurrent for less than 8 mm. Tendrils ±1.5–2 times the length of the lamina, generally 12–20 cmlong, often curling twice. Lower pitcherscylindric to sub-ellipsoid throughout or narrowly 2The emended description is based on measurements made of herbarium specimens and in situmaterial, and is provided in order to supplement the incomplete description presented in Adam & Hamid (2006). ellipsoid in lower half and cylindric above, often narrowing slightly towards mouth, 12–16(–23) cm tall, 3–5 cmwide, with fringed wings 2–3 mmwide, fringe elements to 7 mmlong, 3–6 mmapart; peristome oblique, typically wider towards raised neck, neck often recurved forwards over mouth, 3–5 mmwide at front of pitcher, sub-cylindric, slightly flattened, increasingly so towards neck and broadening to 6–8 mmwide, ribs ca. 0.3–0.4 mmapart, teeth minute, ≤ 0.4 mmlong. Lid ovate to sub-triangular, 3–5 cmlong, 1–1.8 cmwide, margins sinuate, occasionally revolute, abaxial midline rib pronounced, with semi-circular basal keel to 3.5 mmtall, distal part sometimes with apical protuberance 2–3 mmlong, bearing ocellate-crateriform glands 0.5–0.8 mmin diameter, abaxial surface of lid with small, scattered, pitted glands ca. 0.2 mmin diameter.Spur simple, 5–8 mmlong. Upper pitchersgenerally infundibular with a prominent ventral gibbosity below peristome, rarely narrowly infundibular throughout, 10–16(–19) cm tall, 4–6 cmwide, wings reduced to prominent ridges, peristome sub-cylindric, somewhat flattened, 2–6 mmwide at front and ca.1/3 wider at rear, ±horizontal at front, rising sharply towards rear into neck, neck often recurved over mouth. Lid narrowly triangular-ovate, appearing ligulate due to revolute margins, 1.5–3.8(–5) cm long, generally>4.5 times longer than wide, typically 0.5–1 cmwide, abaxial midline rib with pronounced semi-circular basal keel, apex with or without glandular apical protuberance up to 3 mmlong bearing ocellate-crateriform glands, same glands sometimes present along abaxial midline rib, small nectar glands present across abaxial surface. Spur simple, filiform, 4–12 mmlong. Male inflorescenceto ca. 20 cmlong, 30–80 flowers, peduncle 3–8 cmlong, rachis 9–15 cmlong, partial peduncles 2-flowered, pedicels ca. 0.8 mmlong, tepals elliptic, 3–4 mmlong, 2.5–3 mmwide, apex acute, staminal column 3–4 mmlong, anther head 1.5 mmin diameter. Female inflorescenceto ca. 28 cmlong, 30–50 flowers, peduncle 5–12 cmlong, rachis 4–13(–20) cm long, partial peduncles 2-flowered, pedicels 0.8–1.2 cmlong, tepals elliptic in shape, 3.5–5 mmlong, 2.5–3 mmwide, apex acute, fruit 2.5–3 cmlong, seeds filiform, 1.2–1.8 cmlong, pale brown. Indumentumvariable, consisting of short, simple, rufous to white hairs present on stems, pitchers, leaf margins, abaxial leaf surfaces, abaxial midribs, and on developing foliage, becoming caducous in some plants such that older stems and leaves may appear sub-glabrous, generally denser and more persistent on inflorescences. Colourof stems and leaves green, new foliage sometimes suffused with red, pitcher colouration highly variable, lower pitchers green, grey-green or yellow-green, usually mottled red, purple or black, peristome green, yellow to almost black, typically striated with bands of red or purple, but often of uniform colour, upper pitchers equally variable, externally solid yellow-green or with red to purplish-brown blotches, peristome often striated yellow and red, occasionally wholly yellow or red.  Phenology:—Herbarium specimens of  Nepenthes zakrianain flower exist for almost every month of the year, suggesting that flowering may occur at any point in time for a given individual.   Distribution and ecology:—  Nepenthes zakrianahas been recorded with certainty only from Sabah, and chiefly the Crocker Range ( Fig. 4), with Mt. Kinabalu and surrounding areas being its locus classicus. A single, pitcherless specimen from Sarawak’s Gunung Mulu National Park ( (Jermy) 13253) is labelled as  N. fuscaand resembles  N. zakriana(lacking the decurrent leaf bases of  N. dactylifera), but its laminae are broadly elliptic with obtuse apices and abruptly attenuated bases, atypical for the species; more complete material would be needed to confirm the identity of this taxon.   Nepenthes zakrianais generally found at elevations of 1200–2500 m, but occasionally down to  800 m.This species appears to be predominantly epiphytic, but is often observed growing terrestrially above 1300 m, being ruderal at sites like road cuttings and on serpentine rubble, but also occurring on natural humus and moss banks. Like most  Nepenthes,  N. zakrianahybridises readily with its congeners. The literature records putative natural hybrids between  N. zakriana(under the name  N. fusca) and the following species:  N. burbidgeaeHook.f ex Burbidge (1882: 56)on Mt. Kinabalu and adjacent areas such as Mamut copper mine (AR, MG pers. observ., Lowrie 1983);  N. lowiion Mt. Alab ( Phillipps et al.2008; originally identified as  N. lowii×  N. pilosa Danser (1928: 355)by Clarke 1997) and Mt. Trus Madi ( Fretwell 2013);  N. reinwardtiananear Mt. Kinabalu (S. Hirosi pers. comm., McPherson 2009;  N. naquiyuddinii Adam & Hamid (2006: 431)may represent this cross, per Phillipps et al.2008);  N. stenophylla Masters (1890: 240)on Mt. Trus Madi ( Phillipps et al.2008) and Mamut copper mine (AR pers. observ., Thong 2006);  N. tentaculataon Mt. Alab ( Phillipps et al.2008); and  N. rajah Hooker (1859: 421)in the Mt. Kinabalu area ( Clarke 1997).  Conservation status:—  Nepenthes zakrianaoccurs widely across the western highlands of Sabahand is found in a number of protected areas including the Crocker Range NP and Kinabalu NP. It qualifies as Least Concernwhen evaluated against the IUCN 3.1 criteria ( IUCN 2012).   FIGURE 9.  Nepenthes zakriana(J.H.Adam & Wilcock) J.H.Adam & Hafiza. A—climbing stem. B—lower pitcher. C—abaxial surface of lid of lower pitcher. D—upper pitchers, top rightMamut copper mine, bottom left, Mt. Trus Madi. E—abaxial surface of lid of upper pitcher. F—cross section of peristome. G—inflorescence. H—dehiscing seed capsules. Scale bars: A, B, D, G, H = 1 cm, C, E, F = 5 mm. Based on Beaman 9957and Hobbs 17, and supplemented with photographs and measurements made of in situmaterial. Illustrated by M. Barer.   FIGURE 10.Pitchers of  Nepenthes zakriana. A—lower pitcher (Mamut copper mine). B—lower pitcher (Mt. Trus Madi). C—upper pitcher (Mt. Trus Madi). D—upper pitcher (Mt. Alab). E—climbing stem with pitchers, showing non-decurrent leaf bases (Mamut copper mine). Photographs A–E by A.S. Robinson.  Notes:—The name  Nepenthes zakrianais applied to this taxon as it was validly published by Adam & Hamid (2006)to describe typematerial collected from a well-characterised and representative population of  N. fusca sensu Kurata (1976)from Mamut copper mine (Mt. Kinabalu). However, the protuberant apical lid appendage cited by Adam & Hamid (2006)as a key characteristic of  N. zakrianacannot be regarded as a reliable diagnostic feature of this species. Different plants within a single population may or may not produce apical appendages, while different pitchers on a single plant can be equally variable—an inconsistency that can be verified at the typelocality (AR, MG pers. observ.), as well as on trails around the Kinabalu Park Headquarters, Mt. Alab and Mt. Trus Madi (AR pers. observ.). However, other features (listed in Table 2) including leaf base decurrency, lid morphology, pitcher shape and indumentum make it possible to distinguish readily between this taxon and the closely related  N. dactylifera.  Additional specimens examined:—   SABAH:  Beaman8178(K!, L!; photo), Penampang District, Crocker Range, Km50.7 on Kota Kinabalu–Tambunan Road, 5°50’N 116°19’E,  1600 m,  2 January 1984[K, L—climbing stem with upper pitcher];   Beaman8910(K! [2 sheets]), Penampang District, Crocker Range, Km51.8 on Kota Kinabalu– Tambunan Road, 5°50’N 116°20’E,  1500–1600 m,  16 March 1984[separate leaves and pitchers; pitcherless climbing stem];   Beaman9957(K!, L!; photo), Ranau District, Mamut Copper Mine, 6°02’N 116°39’E,  1600–1700 m,  30 May 1984[K—climbing stem with upper pitchers, separate lower pitcher; L—leaf with lower pitcher, separate lower pitchers]; ( Clemens& Clemens)  30980/31092( BO!), single sheet combining two labels: ( 30980) Upper Kinabalu, Penibukan, near Tablerock, left trail, north ridge,  5000 ft[=  1524 m],  16 January 1933; ( 31092)  Mt. Kinabalu, Penibukan, ridge above Kina Taki river,  4000 ft[=  1219 m],  16 January 1933[climbing stem with upper pitchers, infructescence; “  N. stenoperculumMSC”—annotation by M.S. Clemens]; ( Clemens& Clemens)  31570( BO!), two conflicting labels: (1) Mt. Kinabalu, Penibukan,  4000–5000 ft[=  1219–1524 m],  7 February 1933; (2)  Mt. Kinabalu, Keebambang river,  4000 ft[=  1219 m],  3 August 1933[climbing stem with upper pitchers, male inflorescence]; ( Clemens& Clemens)  34454(BO! [2 sheets]), Mt. Kinabalu, Colombonbasin, below Keebambanglobang,  3500 ft[=  1067 m],  14 August 1933[climbing stems with upper pitchers]; ( Clemens& Clemens)  50961(K!), Upper Kinabalu,  7000–8000 ft[=  2134–2438 m],  7 December 1933[climbing stem with upper pitchers];   Collenette815[a](K!), Royal Society Route[up Mt. Kinabalu], near platform 2 hrs above Base,  5275 ft[=  1608 m],  16 August 1961[separate leaf and lower pitcher elements];   Collenette815[b](K!), Mt. Kinabalu, eastern shoulder,  4000–5000 ft[=  1219–1524 m],  16 August 1961[short stem with lower pitchers];   Collenette816[a](K!), Royal Society Route[up Mt. Kinabalu], near platform 2 hrs above Base,  5275 ft[=  1608 m],  16 August 1961[leaf with upper pitcher];   Collenette816[b](K!), Mt. Kinabalu, eastern shoulder,  4000–5000 ft[=  1219–1524 m],  16 August 1961[climbing stem with upper pitchers];   Comber4031(K!), Sapong,  3000 ft[=  914 m], no date [climbing stem with upper pitchers, male inflorescences];   de Vogel8078( L!), West Coast Residency, road Kimanis–Keningau, Crocker Range, on the watershed, 5°28’N 116°03’E,  1500 m,  6 October 1986[stem with intermediate pitchers];   Hobbs17(K!), Trus Madi, SE ridge,  4500 ft[=  1372 m],  22 August 1977[pitcherless climbing stem and leaf with lower pitcher];   Hobbs18(K!), Trus Madi, SE ridge,  4500 ft[=  1372 m],  22 August 1977[short stem with lower pitchers];   Hobbs72(K!), Ranau, Mt. Kinabalu,  5500 ft[=  1676 m],  26 September 1977[climbing stem with intermediate pitcher, separate lower pitcher];   Hobbs73(K!), Ranau, Mt. Kinabalu,  6500 ft[=  1981 m],  26 September 1977[short stem with lower pitchers];   Rickards101(K!), Mt. Kinabalu,  5400 ft[=  1646 m],  19 June 1976[short stem with lower pitchers];   Rickards156(K!), Mt. Kinabalu,  5500 ft[=  1676 m],  20 June 1976[short stem with lower pitchers];  RSNB 4377 ( Chew& Corner)(K!), Bembangan River,  5000 ft[=  1524 m],  19 February 1964[climbing stem with upper pitchers, male inflorescence];  RSNB 4415 ( Chew& Corner)(K!), Bembangan River,  5000 ft[=  1524 m],  20 February 1964[stems with lower pitchers];   SAN 60818( Aminet al.)(K!), Tambunan District, Gu. Alab Range,  600 ft[=  183 m] [ sic—clearly an error; the elevation of Mt. Alab is ca.  6200 ft(  1900 m)],  21 July 1984[climbing stem with upper pitcher, female inflorescence];   SAN 82759( Lantoh)(K!, KEP, L, SAR), Ranau District, Kinabalu Sabah National Park,  6000 ft[=  1829 m],  26 January 1976[K—climbing stem with upper pitchers, male inflorescences];   SAN 91184( Aban& Dewol)(K!, KEP, L, SAR, SING), Beluran District, Bukit Liminintong,  2800 ft[=  853 m],  9 April 1980[K—short stem with lower pitchers];   SAN 91185( Aban& Dewol)(BO, K!, KEP, L, SAR, SING), Beluran District, Bukit Liminintong,  2800 ft[=  853 m],  9 April 1980[K—climbing stem with upper pitcher, infructescence];   SAN 121014( Aminet al.)(K!, L!), Ranau District, Bambangan, no elevation data,  10 September 1987[K, L—climbing stem with intermediate pitcher];   SAN 127727( Fidilis)(K!, L!), Penampang District, Togudon/ Tungol Km48 Jalan Tambunan/ Penampang,  1400 m,  14 August 1989[K—two climbing stems: one with upper pitcher, one pitcherless with male inflorescence; L—pitcherless climbing stem with infructescence];   SAN 147507( Suzana)(K!, L!), Ranau, Mamut Copper Mine, 6°20’N 116°40’E,  1200 m,  29 November 2005[K—climbing stem with upper pitcher, female inflorescence; L—pitcherless climbing stem with female inflorescence];   Schwallier60( L!), Crocker Range, down main road [from] Alab Station, 5°49’4.09”N 116°20’23.16”E,  1836 m,  24 September 2012[climbing stem with upper pitcher, male inflorescence];   Schwallier61( L!), Crocker Range, down main road [from] Alab Station, 5°48’51.2”N 116°20’23.58”E,  1778 m,  24 September 2012[climbing stem with upper pitcher, female inflorescence (unopened flowers)];   Schwallier62( L!), Crocker Range, down main road [from] Alab Station, 5°48’52.38”N 116°20’23.82”E,  1764 m,  24 September 2012[short stem with lower pitcher];   Schwallier63( L!), Crocker Range, down main road [from] Alab Station, 5°48’52.38”N 116°20’23.82”E,  1764 m,  24 September 2012[climbing stem with upper pitcher, inflorescence (unopened flowers)].  SARAWAK: (Jermy) 13253(K!), Gunung Mulu National Park, around Camp 3 [of Royal Geographical Society’s Expedition to Gunong Mulu 1977–78],  1150– 1250 m,  5 October 1976[pitcherless climbing stem—in the absence of pitchers, the specimen is of limited value].   Nepenthes stenophylla:—   SABAH:  SAN 83146(Cockburn)(K!, L), Lamag District, Inarat, S.ridge of Gunung Lotung,  3000 ft[=  914 m],  15 May 1976[K—high epiphyte; climbing stem with upper pitcher, lid absent];   SAN 83251(Cockburn)(K!, L), Lamag District, Inarat, S.slopes of Gunong Lotong[ Maliau Basin],  4300 ft[=  1311 m],  16 May 1976[K—climbing stem with upper pitcher, infructescence];   SAN 151747( Pereira)( L!), Ranau, west of  Bkt. HampuanFR[Forest Reserve], 6°01’26”N116°39’41”,  1489 m,  12 May 2010[climbing stem with upper pitcher, infructescence].  Notes on specimens examined:— SAN 83251 (Cockburn)was labelled as  Nepenthescf. stenophyllabut subsequently identified as  N. fuscaby Smythies ( 20/12/1977) and Jebb ( 9/11/1993) on the sheet. However, the lid, though partially folded, appears complanate and is noted to be as large as the pitcher opening, which is atypical of  N. fusca s.lat.—based on extensive exploration of the Maliau Basin, AR, SM and BQ determine that this material represents the diminutive heath-forest form of  N. stenophyllafound throughout the Maliau Basin above elevations of ca. 1000 m. SAN 83146 (Cockburn)is conspecific with this taxon. SAN 151747 (Pereira), identified as  N. fuscaby Madani (7/2010), likewise represents  N. stenophylla(det. AR & MG), which is widespread at the stated locality.  The isotypeof  Nepenthes zakrianaat the Universityof Aberdeen Herbarium( ABD) could not be located despite an extensive search of the collections; staff indicate that there are no records that it was ever deposited. 1984-01-02 L Beaman Penampang District 1600 5.8333335 Km 1305 116.316666 Crocker Range 20 117 1 Sabah 1984-03-16 Beaman 1550 21 118 5.8333335 Crocker Range 1305 116.333336 Penampang District 20 117 1 Sabah 1984-05-30 L Beaman & Clemens 1650 6.0333333 Mamut Copper Mine 1305 116.65 Ranau District 21 118 1 Sabah 1933-01-16 BO Upper Kinabalu 1524 Table Penibukan 21 118 1 Sabah 1933-01-16 M. S. Clemens & Clemens Mt. Kinabalu 1219 Kina Taki river Penibukan 21 118 1 Sabah 1933-02-07 BO 1372 Penibukan Mt. Kinabalu 21 118 1 Sabah 1933-08-03 Clemens 1219 Keebambang river Mt. Kinabalu 21 118 1 Sabah 1933-08-14 Clemens Mt. Kinabalu 1067 Keebambang Colombon 21 118 1 Sabah 1933-12-07 2286 Upper Kinabalu 21 118 1 Sabah 1961-08-16 Collenette 1608 Base Royal Society Route 21 118 1 Sabah 1961-08-16 1372 Mt. Kinabalu Collenette 21 118 1 Sabah 1961-08-16 Collenette 1608 Base Royal Society Route 21 118 1 Sabah 1961-08-16 1372 Mt. Kinabalu Collenette 21 118 1 Sabah [136,1412,736,761] 914 Sapong Comber 21 118 1 Sabah 1986-10-06 L de Vogel West Coast Residency 1500 5.4666667 Crocker Range 1305 116.05 Kimanis - Keningau 21 118 1 Sabah 1977-08-22 1372 Trus Madi Hobbs 21 118 1 Sabah 1977-08-22 1372 Trus Madi Hobbs 21 118 1 Sabah 1977-09-26 Hobbs 1676 Mt. Kinabalu Ranau 21 118 1 Sabah 1977-09-26 Hobbs 1981 Mt. Kinabalu Ranau 21 118 1 Sabah 1976-06-19 1646 Mt. Kinabalu Rickards 21 118 1 Sabah 1976-06-20 1676 Mt. Kinabalu Rickards 21 118 1 Sabah 1964-02-19 Chew & Corner 1524 Bembangan River 21 118 1 Sabah 1964-02-20 Chew & Corner 1524 Bembangan River 21 118 1 Sabah 1984-07-21 SAN Amin 183 Gu. Alab Range Tambunan District 21 118 SAN 60818 1 Sabah 1976-01-26 KEP, L, SAR Lantoh 1829 Kinabalu Sabah National Park Ranau District 21 118 SAN 82759 1 Sabah 1980-04-09 KEP, L, SAR, SING Aban & Dewol 853 Bukit Liminintong Beluran District 21 118 SAN 91184 1 Sabah 1980-04-09 KEP, L, SAR, SING Aban & Dewol 853 Bukit Liminintong Beluran District 21 118 SAN 91185 1 Sabah 1987-09-10 L Amin Bambangan Ranau District 21 118 SAN 121014 1 Sabah 1989-08-14 L Fidilis Penampang District 1400 Tungol Km Togudon 21 118 SAN 127727 1 Sabah 2005-11-29 L Suzana 1200 6.3333335 Mamut Copper Mine 1304 116.666664 Ranau 21 118 SAN 147507 1 Sabah 2012-09-24 L Schwallier 1836 5.817803 Alab Station 1 116.33977 Crocker Range 21 118 1 Sabah 2012-09-24 L Schwallier 1778 5.8142223 Alab Station 1 116.33989 Crocker Range 21 118 1 Sabah 2012-09-24 L Schwallier 1764 5.8145504 Alab Station 1 116.33995 Crocker Range 21 118 1 Sabah 1976-10-05 2012-09-24 1976-10-05 L Malaysia Schwallier 1764 5.8145504 Alab Station 1 116.33995 Crocker Range 21 118 1 1976-05-15 L Inarat, S. & Gunung Lotung 914 Lamag District 21 118 SAN 83146 1 Sabah 1976-05-16 L Inarat, S. & Gunong Lotong & Maliau Basin 1311 Lamag District 21 118 SAN 83251 1 Sabah 2010-05-12 L Pereira & Ranau & Bkt. Hampuan FR 1489 6.023889 Bkt. Hampuan 21 118 SAN 151747 1 Sabah ABD Herbarium University 22 119 1 Aberdeen City isotype