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Nepenthes erucoides (Nepenthaceae), an ultramaficolous micro-endemic from Dinagat Islands Province, northern Mindanao, Philippines


Abstract and Figures

A new species of Nepenthes—N. erucoides—is described and illustrated from a single ultramafic peak in the Dinagat Islands Province of northeastern Mindanao. It is a distinctive component of a relatively low-elevation, highly biodiverse montane elfin forest that has evolved in association with a particularly thin and extremely hostile substrate. Plant habit, and leaf, inflorescence, indumentum and peristome-column morphology appear superficially similar to those of the ultramaficolous montane species of Palawan, implicating an environmental basis for a syndrome of shared characteristics.
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Phytotaxa 423 (1): 021–032
Copyright © 2019 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Duilio Iamonico: 21 Oct. 2019; published: 31 Oct. 2019
Licensed under a Creative Commons Attribution License
Nepenthes erucoides (Nepenthaceae), an ultramaficolous micro-endemic from
Dinagat Islands Province, northern Mindanao, Philippines
1 National Herbarium of Victoria, Royal Botanic Gardens Melbourne, Melbourne, VIC 3004, Australia;
2 The Graduate School, University of Santo Tomas, España Boulevard, 1015 Manila, Philippines.
3 Research Center for the Natural and Applied Sciences, University of Santo Tomas, España Boulevard, 1015 Manila, Philippines.
4 DENR-PENRO Dinagat Islands, Caraga Region Purok 2, Barangay Santa Cruz, San Jose, Province of Dinagat Islands, Mindanao,
*Author for correspondence
A new species of NepenthesN. erucoides—is described and illustrated from a single ultramafic peak in the Dinagat Islands
Province of northeastern Mindanao. It is a distinctive component of a relatively low-elevation, highly biodiverse montane
elfin forest that has evolved in association with a particularly thin and extremely hostile substrate. Plant habit, and leaf,
inflorescence, indumentum and peristome-column morphology appear superficially similar to those of the ultramaficolous
montane species of Palawan, implicating an environmental basis for a syndrome of shared characteristics.
Keywords: Philippines, Malesia, new species, non-core Caryophyllales, taxonomy, ultramafic
Nepenthes Linnaeus (1753b: 955) is a genus of tropical pitcher plants comprising ca. 160 species (see e.g., Clarke
et al. 2018), with a wide distribution across the Malesian and Papuasian biogeographic regions, with recognised
centres of diversity in Borneo, the Philippines and Sumatra, where the greater proportion of species are represented
by microendemic taxa from individual mountains or mountain ranges (Cheek & Jebb 2001, Clarke 2001, Robinson et
al. 2009).
In 2001, only 12 species of Nepenthes were recognised from the Philippines (Cheek & Jebb 2013), but 44 further
species have been described from the country since then. This marked increase stems from a number of factors, including
novel discoveries resulting from increased exploration of the archipelago by botanists and Nepenthes enthusiasts alike;
a concomitant increase in familiarity with the genus amongst Filipino botanists and, particularly, a Kew-led project that
has elevated a raft of taxa to species rank, mainly from overlooked specimens in herbaria around the world, and raised
others from synonymy (e.g., Cheek & Jebb 2014, 2013a, 2013b, 2013c, 2013d).
When excluding those newly described Filipino species that are not accepted by various authors (e.g., Clarke et al.
2018), the total number of good species recognised from the archipelago still stands at approximately 45. Out of the 56
taxa described at specific rank, ca. 30 are either ultramafic endemics or largely restricted to mafic substrates or surface
laterites with associated open canopies. A further ca. 20 taxa are associated with more general mid-montane and
montane habitats, mainly in association with mossy forest where they predominantly grow as epiphytes or in pockets of
humus. Though many in this group are recorded from volcanoes, few Filipino Nepenthes are actually recorded growing
directly in volcanic substrates, rather occurring in humic material overlying volcanic substrata. In fact, the only Filipino
Nepenthes verifiably documented as facultatively growing directly in volcanic substrates are N. ventricosa Blanco
(1837: 807), e.g. on the lava tube walls of the Mayon volcano, and N. copelandii Merrill ex Macfarlane (1908: 51), e.g.
on volcanic rubble on the slopes of the Camiguin and Mt. Apo volcanoes (A. Robinson, pers. observ.). The few other
non-ultramaficolous terrestrial species known are typically recorded from other substrates, such as weathered granites
[e.g., N. mira Jebb & Cheek (1998: 966), Cleopatra’s Needle], degraded shale and mudstone [e.g., N. robcantleyi
Cheek (2011: 678) at one of two mountain locations above the Compostela Valley, also epiphytic at lower elevations]
22 Phytotaxa 423 (1) © 2019 Magnolia Press
and limestone [e.g., N. viridis Micheler et al. (2012: 4), coastal Dinagat] (all A. Robinson, pers. observ., but see also
Encarnación & Mukasa 1997, and Suerte et al. 2009).
Within the Philippines, the highest Nepenthes species diversity is recorded on Mindanao, where 30 species occur
and 18 of them are ultramaficolous. This high species diversity correlates with the spatial heterogeneity of exposed
ultramafic terranes on Mindanao, which include numerous isolated mountain peaks. The edaphic effects of nutrient-
poor, broadly phytotoxic, cation-imbalanced and frequently water-stressed soils combined with the particular climatic
factors associated with individual ultramafic outcrops are disproportionately associated with high levels of speciation
and endemism (Galey et al. 2017). The eastern half of Mindanao is home to the largest tracts of exposed ultramafic
regolith on that island, and the prominent outcrops of Surigao del Norte Province, Dinagat Islands Province and
the Pujada Peninsula (Davao Oriental, including Mt. Hamiguitan) are all derived from the same belt of Cretaceous
ophiolite complexes (Balce et al. 1976, Yumul et al. 2003, 2008, Tamayo et al. 2004), a geological feature associated
with a range of endemic Nepenthes including N. bellii Kondo (1969: 653), N. hamiguitanensis Gronemeyer et al.
(2010: 1296), N. merrilliana Macfarlane (1911: 207), N. micramphora Heinrich et al. (2009: 1315), N. mindanaoensis
Kurata (2001: 32), N. peltata Kurata (2008: 12), N. ramos Jebb & Cheek (in Cheek & Jebb 2013d: 108), N. surigaoensis
Elmer (1915: 2785) and N. truncata Macfarlane (1911: 209). Six Nepenthes species, i.e. N. bellii, N. merrilliana,
N. mindanaoensis, N. mirabilis (Loureiro 1790: 606) Rafarin (1869: 270), N. truncata, and N. viridis (only on
limestone within this region), have been recorded just from the fraction of the relatively small Dinagat Islands Province
(ca. 1036 km2) already investigated by the first author.
In November 2014, an unidentified species of Nepenthes was photographed from a conserved, relatively low-
elevation ‘montane forest’ on Dinagat Island by botanists J. Barcelona and P. Pelser (University of Canterbury, New
Zealand) as part of their ongoing work to catalogue the entire flora of the Philippines (Pelser et al. 2011+). Images
of the plant were shared with AR, who confirmed its undescribed status. Assessments of the plant were carried out in
2018, after which representative material was collected and accessioned, allowing for its description, presented here.
Materials and methods
In situ studies of Nepenthes were made on Dinagat by AR in July 2014, and specific morphological and ecological
studies of N. erucoides and its habitat were conducted by AR and Brian Quinn (New Zealand) with the support of RC
in June 2018 after a permit for access to the habitat was granted. Material of the plant was collected by SZ (under
DENR Region XIII Wildlife Gratuitous Permit No. R13-2018-0053 and Wildlife Transport Permit No. 2018-10-01)
and subsequently deposited at the herbaria PNH and UST (acronyms follow Thiers 2019+). Fine measurements were
made using a Mitutoyo vernier calliper (Mitutoyo Corporation, Japan).
Initial de novo georeferencing was made using a Garmin Oregon 600 GPS unit with dual GPS and GLONASS
telemetry enabled. Key measurements were made with 3 averaged waypoint readings over the course of a 15 minute
period, with an estimated accuracy of ±3 m.
Nepenthes erucoides A.S.Rob. & S.G.Zamudio, sp. nov. (Figs 1–3).
Type:—PHILIPPINES. Mindanao: Dinagat Province, Mt. Redondo, 839 m elevation, 13 October 2018, Zamudio, Apo, Gonzales &
Tan 257395 (holotype, PNH!) [mature rosette bearing three lower pitchers]; ibidem Zamudio, Apo, Gonzales & Tan 014747 [detached
upper pitcher], 014749 [three sheets: mature rosette with tendrils; lower pitcher × 2; upper pitcher], 014753 [intermediate pitcher]
and 014754 [small rosette with very dense indumentum] (all isotypes, UST!).
Diagnosis:Nepenthes erucoides is similar to N. mantalingajanensis Nerz & Wistuba (2007: 17) in overall plant
form, but differs from that species by way of its (differences in parentheses) long, dense indumentum of rufous hairs
(predominantly glabrous), production of lower and upper pitchers (lower pitchers only), urceolate to sub-campanulate
pitcher shape (ovate-obconic), finer peristome with small teeth along the inner margin (broader with clearly defined
ribs and long teeth), very large bracts 8–15 mm long produced from some partial peduncles and the rachis of male
inflorescences (bracts on most partial peduncles, ca. 1 mm long, never from the rachis), and seeds without well-
developed filaments (filaments well-developed).
THE MICRO-ENDEMIC NEPENTHES ERUCOIDES Phytotaxa 423 (1) © 2019 Magnolia Press 23
FIGURE 1. Nepenthes erucoides. A) mature rosette bearing female inflorescence and upper pitcher; the prominent, dense, adaxial
indumentum of the newest leaf soon becomes caducous. B) lower pitcher. C) upper pitcher in section; the waxy zone is present only on the
column and just below the peristome. D) adaxial (left) and abaxial (right) surfaces of a pitcher lid. E) detail of a newly formed leaf with
its dense indumentum and a developing leaf; the stem indumentum is variably caducous between individuals. F) entire (left) and sectioned
(right) indehiscent fruit. G) male flower with very large bract. H) adaxial (top), lateral section (middle) and abaxial (bottom) detail of tepal
from male flower. Scale bars: A, B, C, D, E = 1 cm, F = 5 mm, G, H = 1mm. Based on Zamudio, Apo, Gonzales & Tan 257395 and 014754,
as well as on photographs and measurements made in situ. Illustrated by A. Robinson.
24 Phytotaxa 423 (1) © 2019 Magnolia Press
FIGURE 2. A) mature rosette emergent from elfin vegetation. B) lower pitcher. C) upper pitcher. D) the limited waxy zone. E) transverse
sections of (left) lower and (right) upper pitchers showing the almost entirely glandular interior. F) detail of the superficially caterpillar-
like (erucoid) developing leaf emergent from the petiolar-laminar groove of the preceding leaf. G) a recently unfurled leaf; the dense
adaxial indumentum has already begun to be shed towards the leaf base. Photographs A, C by P. Pelser; B, D–G by A. Robinson.
THE MICRO-ENDEMIC NEPENTHES ERUCOIDES Phytotaxa 423 (1) © 2019 Magnolia Press 25
FIGURE 3. A) the indumentum of juvenile stem and leaf parts is abruptly caducous. B) the largely persistent indumentum of the tendril.
C) the extremely dense indumentum of a senescent male inflorescence; note the large bracts on the rachis and partial peduncles. D)
sequential anthesis and development of tepal colouration in the male inflorescence. E) detail of male flowers at anthesis. F) a female
rosette in fruit. G) transverse section of fruit with seeds; note the relatively short, broad form of the seeds. H) the natural hybrid Nepenthes
erucoides × mindanaoensis. I) the elfin ‘forest’ at the summit of Mt. Redondo, formed in response to the inorganic, granular, ultramafic
substrate and associated conditions. Photograph A by P. Pelser; B–H by A. Robinson.
26 Phytotaxa 423 (1) © 2019 Magnolia Press
Description1:—An upright to subscandent sub-shrub 10–50(–120) cm tall. Stems terete, 0.9–1.8 cm in diameter,
internodes 0.5–3 cm long, climbing stems absent. Leaves coriaceous, petiolate, emerging pliant, becoming brittle on
maturity, lamina broadly sub-elliptic to oblong, 8–15 cm long, 4.5–8.5 cm wide, apex retuse-emarginate and often
unequal either side of midrib, base of lamina abruptly attenuate into petiole, petioles 0.8–1.8 cm long initially, but
up to 3.8 cm long on taller stems, without wings, base sheathing the stem and amplexicaul for 1/2–4/5 of the stem
circumference, initially tubular, wholly enclosing primordium of subsequent leaf, opening to form a canaliculate
petiolar-laminar groove. Longitudinal veins 2(–3), more conspicuous on abaxial surface, situated on either side of
midrib in outer 1/3 of lamina, parallel to margin, with fine, ±simple transverse venation running between longitudinal
veins and midrib. Tendrils rarely peltate from apex of large leaves, ca. 2.5 times longer than the laminae, 25–38 cm
long, uncoiled in lower and upper pitchers. Lower pitchers urceolate to broadly sub-cylindric or sub-campanulate,
6–10(–15) cm tall, 4.5–8.5 cm wide, typically widest at the mouth, tendril ventrally attached, ventral surface between
wings flattened, largest pitchers becoming angular, wings poorly developed, present only in the most basal lower
pitchers, thereafter reduced to pronounced ridges, restricted to upper 1/3 of ascidium if manifested at all, and 1–4 mm
wide, fringe elements absent or sparse and tubercle-like, 0.5–1.2 mm long, 4–6 mm apart; pitcher interior almost
entirely glandular, bar a scant waxy zone arising only near the base of the column and continuing to its apex, also
contiguous but hidden beneath inner margin of peristome, glands in lower 1/6 of pitcher slate grey to brown in colour,
elliptic, 0.3–0.5 mm long, 0.2–0.3 mm wide, with a density of ca. 330 glands per cm2, glands in upper 5/6 of pitcher
of same colour as pitcher interior, 1/3 smaller and more densely arranged, with a density of ca. 400 glands per cm2;
peristome slightly raised to ±horizontal at front of pitcher between wings, thereafter rising at an angle of 20°–25°
towards pitcher rear, becoming abruptly vertical at column, column itself typically recurved forwards over pitcher
opening at an angle of ca. 30° from vertical, peristome in section moderately curved to sub-cylindric at the front,
2–4(–6) mm wide, rear part broadening only slightly by less than 1/3, becoming flattened and appressed from the base
of the column to its apex, external marginal lobing absent, ribs fine but pronounced, ca. 0.2–0.4 mm high, ca. 0.8 mm
apart, teeth triangular-acuminate, 0.8–1 mm long along inner margin. Lid positioned between horizontal and 45°,
cordate, sometimes ±complanate in smaller pitchers, but typically broadly canaliculate along the midvein, canaliculum
occupying central 1/3 of lid between a pair of perfect or imperfect, basal to supra-basal acrodromous veins, one
each side of the canaliculum furrow, apex retuse to obtuse, 2.5–5 cm long, 3.2–5.8 cm wide, abaxial surface without
midline rib protuberances, but densely, uniformly covered with minute, elliptic to round pitted glands. Spur vestigial
to simple and short, 1–2.8(–5) mm long, rarely bifurcating. Upper pitchers globose to sub-campanulate, attenuating
into the dorsi-basally attached tendril in the lower 1/6, 4.5–9(–11) cm tall, 4–8 cm wide, usually widest just above the
midsection, ventral surface gibbous to slightly flattened, wings usually reduced to ridges, rarely expressed towards
peristome, if so 0.5–3 mm wide, fringe elements usually absent; pitcher interior almost entirely glandular, scant waxy
zone arising only near the base of the column and continuing to its apex as well as immediately beneath teeth of
peristome, glands in lower 2/5 of pitcher slate grey to brown in colour, elliptic, 0.7 mm long, 0.3–0.45 mm wide, with
a density of ca. 165 glands per cm2, glands in upper 3/5 of pitcher of same colour as pitcher interior, 1/3 smaller and
more densely arranged, with a density of ca. 320 glands per cm2; peristome ±horizontal at front of pitcher between
wings, thereafter rising at an angle of 15–20(–40)° towards pitcher rear, becoming abruptly vertical at column, column
itself typically recurved forwards over pitcher opening at an angle of ca. 30° from vertical, peristome in section
sub-cylindric at the front, ca. 2–3.5 mm wide, rear part broadening by up to 1/2, otherwise similar to lower pitchers.
Lid similar to lower pitchers but often slightly shorter and broader, (3–)4.2–5 cm long, (3.5–)4.8–5.8 cm wide. Spur
vestigial, simple, 0.2–1.2 mm long. Inflorescences marcescent, ±emergent from axil at base of preceding leaf. Male
inflorescence 45–55 cm long, 60–80 flowers, peduncle 30–40 cm long, ca. 5–6 mm in diameter at the base, rachis 15–
20 cm long, partial-peduncles 1-flowered, stout, to 1.2 mm in diameter, pedicels 7–9 mm in length, bracts prominent,
simple, terete, ca. 0.35 mm in diameter, 8–15 mm long, produced from underside of only lowermost pedicels or
emergent directly from rachis, adnate to stem, tepals opening yellowish green on the adaxial surface, rapidly turning
deep red to blackish maroon, broadly elliptic, ca. 4.5 mm long, 3 mm wide, concave throughout, adaxial surface with
30–70 elliptic pitted glands, those closest to the base larger than the more distal glands, to 0.4 mm long and 0.18 mm
wide, tepal apex acute; staminal column 2.4–2.8 mm long, anther head ca. 2.3 mm in diameter, anthers ±14, bilocular,
extrorse, thecae pale yellow, pollen grains deep yellow. Female inflorescence ca. 70 cm long, 20–28 flowers, peduncle
55 cm long, 5–6 mm in diameter at the base, rachis 15–18 cm long, partial-peduncles 1-flowered, bracts infrequent
on lowermost pedicels, filiform-subulate, 4–7 mm long, pedicels to 1 mm in diameter, 4–6 mm long, flowers with
a scent of murine urine, fruit 17–20 mm long, 4–5 mm in diameter, tepals as per male flowers but apices acute to
1 This description incorporates details from observations made of living material in situ on Mt. Redondo, as well as from
the holotype (Zamudio, Apo, Gonzales & Tan 257395) and the associated listed isotype materials.
THE MICRO-ENDEMIC NEPENTHES ERUCOIDES Phytotaxa 423 (1) © 2019 Magnolia Press 27
obtuse, glands restricted to central part of adaxial surface, and adaxial surface yellowish green in the proximal 1/3,
and reddish ageing to grey-black in the distal 2/3. Seeds fusiform, ca. 5 mm long, without well-developed filaments.
Indumentum of dense, villous, rufous hairs 2–6 mm long covering all terminal developing parts, tomentose along leaf
margins and abaxial midrib, with downy, rufous to straw-coloured or white hairs 2.5–3 mm long additionally covering
adaxial surface of lamina of most recent leaf; indumentum thereafter abruptly caducous, preceding foliage being
adaxially glabrous and stems frequently so, with indumentum persistent on abaxial surface of lamina and midrib, on
tendril, and more sparsely so on external surface of pitchers, where it is most persistent towards dorsal midrib, as well
as around spur and proximal adaxial surface of lid, with additional dark, minute, stiff hairs contributing to a slightly
scabrous pitcher texture. Indumentum of abaxial lamina surface differs in being predominantly covered with brown,
bristly, curved hairs ca. 2 mm long. Inflorescence indumentum is persistent along the rachis, where it is especially long
and dense in males versus slightly more sparse in females, and close to the base of the peduncle, the peduncle itself
becoming variably sub-glabrous, partial-peduncles with longer hairs on the abaxial surface. The abaxial surface of the
tepals is particularly hairy close to its base and along the midline. Colour of mature stems and leaves a deep, glossy
green, emerging leaves yellowish green, pitchers yellowish olive-green to yellow or orange, sometimes suffused with
red, with or without highly variable amounts of red stippling, peristome colour generally congruent with base colour
of pitcher, adaxial surface of lid similar, but abaxial surface invariable stippled with red, particularly towards base and
Etymology:—The specific epithet erucoides is derived from the Latin eruca (caterpillar) and the Greek suffix –
oides (resembling), in reference to the densely hairy developing leaves which, when still appressed within the petiolar-
laminar groove of the preceding leaf, resemble the exuberantly hairy caterpillars of certain erebid macromoths from
the subfamily Arctiinae, such as those of the genus Arctia Schrank (1802: 152) [e.g. Arctia opulenta Edwards (1881:
Phenology:—Anthesis in Nepenthes erucoides has been observed in June (AR, RC & Brian Quinn, pers. observ.)
and as late as November (Pieter Pelser & Julie Barcelona, pers. comms.), but patterns of inflorescence development
and senescence suggest that the majority of inflorescences emerge in response to increases in rainfall immediately
following the February to April dry season, with peak flowering from late May to early July, and maturation of seed
from late July to early September, immediately preceding the height of the October to January wet season (climate data
from 2019), favouring establishment of seedlings.
Distribution and ecology:Nepenthes erucoides is endemic to Dinagat Island’s highest peak, Mt. Redondo,
where it occurs from ca. 800 m elevation to the 929 m summit in a dark, materially dense, inorganic, granular, clinker-
like substrate of lateritic nickel ore and decomposed chromite rubble, through to a clay derived thereof, certainly the
most extreme ultramafic substrate observed by the first author for any Nepenthes across the entire range of the genus.
This species is a notable component of the extremely stunted [25–70(–150) cm tall] elfin (‘bonsai’) forest that covers
the upper reaches of Mt. Redondo from about 750 m elevation. The forest, which occupies an estimated area of ca. 5.2
km2, is characterised by a shrub-like covering of stunted trees ranging in height from 3 m, at its lowest elevations, to
less than 50 cm in the summit region. Notable plants observed in this region included Dacrydium beccarii Parlatore
(1869: 494) (Podocarpaceae Endl.), Fagraea gitingensis Elmer (1910: 859) (Gentianaceae Juss.), Leptospermum
amboinense Blume (1826: 1100) (Myrtaceae Juss.), Psychotria surigaoensis Sohmer & Davis (2007: 100) (Rubiaceae
Juss.), Rhodomyrtus surigaoensis Elmer (1914: 2344) (Myrtaceae), Scaevola micrantha Presl (1831: 58) (Goodeniaceae
R.Br.), Elaeocarpus Linnaeus (1753a: 515) spp. (Elaeocarpaceae Juss.), several species of Syzygium (Myrtaceae), and
a range of herbaceous plants including Paphiopedilum ciliolare (Reichenback filius 1882: 488) Stein (1892: 462) and
miniature Dendrochilum Blume (1825: 398) spp. (both Orchidaceae Juss.), Dischidia major (Vahl 1810: 110) Merrill
(1917: 437) (Apocynaceae Juss.), Freycinetia Gaudichaud-Beaupré (1824: 509) (Pandanaceae R.Br.) and, of course,
other Nepenthes species, most notably the co-localising N. bellii (where protected from desiccation by thick vegetation)
and N. mindanaoensis (which also hybridises with N. erucoides). N. truncata also occurs in the bonsai forest; however,
within the constraints of the limited observations made during our field studies, this giant species seems to occupy
an adjacent but distinct elevational band mainly below 800 m where the soil profile is deeper and water-holding
capacity greater—N. erucoides is absent from this lower elevation zone. N. merrilliana, which frequently approaches
N. truncata in terms of sheer pitcher size, occurs almost entirely below 600 m elevation in this region.
Within its habitat, Nepenthes erucoides occurs mainly in very open, low growing (<50 cm), scrubby vegetation,
where it is most easily located via its emergent inflorescences. In this situation, shelter from the extremes of intense
sunlight, heat and high rates of evapotranspiration experienced in adjacent open ground likely favours the recruitment
of seedlings. However, scattered plants were also observed growing in apparent good health on rocky, open ground that
was too hot to touch beneath the midday sun, while a minority of plants occurred in denser thickets of taller vegetation
28 Phytotaxa 423 (1) © 2019 Magnolia Press
(up to 1.7 m) where the diminutive N. bellii was occasionally found, if far more difficult to discern. These represented
the tallest and most robust N. erucoides plants (up to 120 cm), however seedling recruitment appeared to be reduced
by low levels of light at ground level as a result of the higher vegetation density.
Conservation Status:Nepenthes erucoides has been directly impacted by mining activities which have resulted
in clearing of part of the Mt. Redondo bonsai forest habitat. However, as part of this work, the mining corporation
overseeing the site (Krominco, Inc.) have set aside a portion of this very important and highly biodiverse habitat for
preservation. The corporation currently acts as conservator of the habitat, critically preventing further damage to the
site and reducing the likelihood of biopiracy by unscrupulous plant and animal collectors by limiting general access
only to permitted and supervised groups.
Direct observations of Nepenthes erucoides made in situ satisfy the IUCN 3.1 Red List CR (Critically Endangered)
Criteria B1ab(ii,iii)+2ab(i,ii,iii) (IUCN, 2012), i.e. the species has an EOO < 100 km2 and is known from only a single
location with documented decline in area of occupancy and quality of habitat, and an AOO < 10 km2 at the single
location with a reduction in area of occurrence, occupancy, and extent and quality of habitat.
Notes:—Although partly caducous, the indumentum of Nepenthes erucoides is the most highly developed in the
genus, being remarkably long and dense on all parts of any primordia, across the entire adaxial surface of the most
recent leaf, and persistent on all floral parts, but particularly the male rachis. This adaptation is presumed to both
protect exposed or developing organs from intense solar radiation and to reduce water loss, particularly during the
very hot dry season (see discussion below). The production of seeds without developed filaments is presumed to be an
adaptation to the species’ island habitat, similar to that seen in N. argentii Jebb & Cheek (1997: 19), N. kerrii M.Catal.
& Kruetr. in Catalano (2010: 30) and N. pervillei Blume (1852: 10); the lack of prominent seed filaments presumably
serves to prevent the dispersal of seeds away from the extremely limited summit elfin forest habitat by strong winds
(A. Robinson, pers. observ.).
Nepenthes erucoides co-occurs with N. mindanaoensis, which also has markedly petiolate leaves, but that species
differs in having a narrower peristome with finer ribs, far less discernible teeth and no column, an abaxially ribbed
lid, as well as 2-flowered partial peduncles and a poorly developed indumentum. Also present, albeit at a far lower
density, is N. bellii, the diminutive sister species to N. merrilliana (which is virtually identical in pitcher, leaf and
to some extent floral characteristics, but differs in its overall miniature form and more sheltered forest ecology), to
which N. erucoides is similar in terms of its lid and the structure of the peristome ribs, but N. bellii has no peristome
column, sessile leaves, and only a sparse indumentum on developing foliage. There are no significant indications
that N. erucoides has a hybrid origin involving these or any of the other species present on Dinagat, and the hybrids
observed between it and N. mindanaoensis show unmistakably intermediate foliar and floral characteristics (Fig. 3H)
as well as, in some cases, extreme hybrid vigour.
The precise nature of the relationship of Nepenthes erucoides with its other congeners is likewise unclear.
In this work it is compared with N. mantalingajanensis based on striking but presumably superficial similarities,
particularly its mostly dwarf, upright habit, its 1-flowered inflorescences, petiolate leaves, the absence of any keel
or processes on the abaxial surface of the lid, and its robust, sometimes markedly-angular pitchers. In this respect,
N. erucoides is similar to another Mindanao Nepenthes, N. peltata of Mt. Hamiguitan, in sharing many characteristics
in common with the ultramaficolous montane species of Palawan and Borneo but differing in ways that cannot easily
be reconciled. The authors are obliged to contend that certain environmental selection pressures common to extreme
ultramafic habitats (e.g., rhizospheric water stresses of largely inorganic substrates, associated hyper-concentrations of
phytotoxic soil cations, exposed habitat-associated transpiration and solar radiation stresses, pronounced temperature
fluctuations even at high elevations) may favour the emergence of a fairly uniform syndrome of robust, ultramafic-
associated morphological characteristics in Nepenthes that confound accurate placement of all of these species based
on morphology alone, highlighting the great need for improved resolution in the markers used to generate meaningful
Nepenthes phylogenies.
Historically, these phylogenies have been poor, however, a recently published analysis using genome skimming has
provided dramatically improved and highly credible phylogenetic and temporal resolution in Nepenthes (Nauheimer
et al. 2019), with the clear potential to address both this suggestion and to more accurately represent the relationships
between members of the genus. A recent preprint (Murphy et al. 2019) also places the robust, morphologically
similar N. mantalingajanensis, N. attenboroughii Robinson et al. (2009: 196), N. deaniana Macfarlane (1908: 57),
N. leonardoi McPherson et al. (2011: 5) and N. palawanensis McPherson et al. (2010: 1332)—all strictly endemic to
the very summits of ultramafic Palawan mountains subject to annual fluctuations in rainfall—as sister to the slender-
pitchered, scrub- to forest-dwelling N. philippinensis Macfarlane (1908: 43), itself sister to N. alata Blanco (1837: 805);
it also places the robust Mindanaoan ultramafic endemic N. peltata, which also resembles the aforementioned Palawan
THE MICRO-ENDEMIC NEPENTHES ERUCOIDES Phytotaxa 423 (1) © 2019 Magnolia Press 29
ultramafic species, in a different clade close to the slender-pitchered N. justinae Gronemeyer et al. (2016: 6) and
N. micramphora. These data seem to support the notion that this syndrome of morphological characteristics may have
a higher likelihood of developing in response to the common environmental challenges of specific ultramafic habitats
irrespective of the clade within which a given species may fall.
The particularly low-elevation elfin forest of Mt. Redondo represents an unusual and especially extreme example
of an ultramafic habitat, the rigours of which may account for the development of this species’ compact and extremely
hirsute characteristics. The occurrence of so extremely stunted a forest at such low elevation cannot be entirely attributed
to the Massenerhebung (mountain mass) effect (Schroter 1926), the elfin forest of Mt. Redondo differing markedly
from the typically wet, lower elevation mossy forests that result from the lowered cloud base that this phenomenon
typically brings about (e.g. Gunung Santubong, Sarawak), particularly in terms of its distinctly sclerophyllous character
(Fig. 3I). Rather, the elfin forest is likely to have formed in response to a range of factors, namely: the geology of Mt.
Redondo, the main prominence of which comprises an 87 million year old exposed ultramafic terrane that affords an
extremely thin, hostile surface substrate of exposed podiform chromite and the unconsolidated weathering products
of the olivine-rich tectonitic rock dunite (Dickey 1975, Madrona et al. 1980, David 1994); seasonal rainfall patterns,
which include a pronounced dry season with average rainfall of ca. 130 mm and particularly hot days of average
ca. 27 °C (see 2019), the effects of which are compounded by the low water-holding capacity of the
almost entirely inorganic soil; and finally by the phytotoxic nature of the soil itself.
The stunting of forests in response to periodic water shortage, but particularly in shallow and stony soils in
more windy and exposed montane situations, is well documented (Van Steenis 1972, Whitmore 1984, Proctor et al.
1988, 1999). The Mt. Redondo elfin forest is exemplary in this regard; the ca. 30 cm deep rhizospheric soil contains,
on average, ca. 396,024 ppm iron (Fe), 1,344 ppm nickel (Ni), 425 ppm copper (Cu), 10,875 ppm chromium (Cr),
and 4,453 ppm manganese (Mn) (Fernando et al. 2018), and 94% of the trees have a stem diameter of just 1–10 cm
(Fernando et al. 2017). Given the rocky, granular nature of ultramafic soils, morphological adaptations to minimise
water requirements and water loss—including reduced leaf morphologies, sclerophyllous characteristics, hirsuteness
and small stature—are particularly well-documented amongst ultramafic-growing plants (Brady et al. 2005); like
the unusual, ultramafic-growing high-montane species N. argentii, N. erucoides appears to represent an extreme,
ultramafic-adapted Nepenthes, albeit one that is specialised to tolerate the added rigours of periodic water stress,
marked temperature fluctuations and high levels of solar radiation, all of which are associated with the low levels of
cloud cover resulting from its windy, relatively low elevation near-coastal habitat.
We thank the Department of Environment and Natural Resources (DENR)—Provincial Environment and Natural
Resources Office (PENRO), Province of Dinagat Islands, especially Mr Leo Joseph E. Oconer, for arranging access
permits and providing key logistical support; Mayor Erwin San Juan and Vice Mayor Pretsy Semorlan of Loreto for
facilitating and seeking to understand the value of our research; and Krominco Inc., particularly staff Gay Espinal and
Rove Cagas, for considering and granting access to the protected forest area within the Mt. Redondo chromite mining
concession. Permission to collect plant specimens from Dinagat Island for scientific study was covered by Gratuitous
Permit No. R13-2018-0053 issued by the Department of Environment and Natural Resources (DENR), Region 13,
CARAGA Regional Office, Butuan City, Philippines. Finally, we thank Julie Barcelona and Pieter Pelser (University
of Canterbury, New Zealand), both for providing data specifically about this new species and for accessory data
supplied via the Co’s Digital Flora of the Philippines database (, Brian Quinn for assistance
with field measurements and logistics, and the Philippine Taxonomic Initiative, Inc. ( for bringing together
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... For example, a survey of the mites collected from six species of Nepenthes in Borneo revealed not only the presence of the inquiline genera Zwickia (two species), Creutzeria (one species), Nepenthacarus (one species) and Naiadacarus (one species), but also two undescribed genera, one containing three species and the other two species. With more than 160 Nepenthes species described to date (Clarke et al., 2018) and a species discovery rate that does not appear to be slowing down (Robinson et al., 2019a(Robinson et al., , 2019b, it is clear that many nepenthebiont mite taxa still await discovery. ...
... In this sense it macroscopically approximates the dense adaxial laminar indumentum of N. erucoides A.S.Rob. & S.Zamudio in Robinson et al. (2019), though the latter is largely caducous and does not take the same form on the pitchers or abaxial surfaces of the leaves. ...
Nepenthes diabolica, a new species of tropical pitcher plant from Central Sulawesi, is described and illustrated. It is a high-montane, terrestrial to epiphytic taxon, distinguished by its elaborately flanged peristome and rufous, woolly pitcher indumentum. N. diabolica is a member of section Tentaculatae, showing morphological affinities with N. hamata. Nevertheless, the two species can be distinguished by morphological characters as well as by ecology and geography. A diagnostic key to the species of section Tentaculatae is provided. Owing to its highly restricted range, small number of recorded individuals, and desirability among plant collectors, the conservation status of N. diabolica is assessed as Critically Endangered under the IUCN criteria. Finally, N. hamata, until now based on lost types and mixed paratypes, is lectotypified using recently uncovered original material.
... This scheme comprises three hierarchical levels of threats increasing in specificity with each level, beginning with 11 first level categories: (1) Residential and Commercial Development, (2) Agriculture and Aquaculture, (3) Energy Production and Mining, (4) Transportation and Service Corridors, (5) Biological Resource Use, (6) Human Intrusions and Disturbance, (7) Natural Systems Modifications, (8) Invasive and Other Problematic Species and Genes, (9) Pollution, (10) Geological Events, and (11) Climate Change and Severe Weather. While second and third level categories further defining threatening processes are sometimes presented in conservation assessments (e.g., Cross, 2020b;Fleischmann et al., 2011Fleischmann et al., , 2017Fleischmann et al., , 2020Gonella et al., 2016;Robinson et al., 2009Robinson et al., , 2018Robinson et al., , 2019b, for the purposes of consistency and statistical analyses we present only first level threatening process categories for species in this study. Data on threatening processes for species assessed by the IUCN were updated where the published literature provided more contemporary evidence. ...
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Carnivorous plants (CPs)—those possessing specific strategies to attract, capture and kill animal prey and obtain nutrition through the absorption of their biomass—are harbingers of anthropogenic degradation and destruction of ecosystems. CPs exhibit highly specialised and often very sensitive ecologies, being generally restricted to nutrient-impoverished habitats where carnivory offers a competitive advantage. As such, they are often the first species to disappear following habitat degradation, land use change, and alteration to natural ecological processes, and are at significant risk from processes such as eutrophication and weed invasion, and even poorly-understood impacts such as airborne nitrogen inputs. Many of the world’s 860 species of CPs are found in wetland habitats, which represent some of the most cleared and heavily degraded ecosystems on Earth. Global diversity hotspots for CPs are likewise located in some of the most heavily cleared and disturbed areas of the planet—southwestern Western Australia, Southeast Asia, Mediterranean Europe, central eastern Brazil, and the southeastern United States—placing their conservation at odds with human developmental interests. Many carnivorous plant species exhibit extreme range-restriction and are wholly localised to specific geological formations, microhabitats or elevations, with nowhere to move to in the face of environmental change such as a warming, drying climate. We provide the first systematic examination of the conservation status and threats to all CPs globally, compiling full or partial assessments of conservation status category for 860 species from 18 genera, and provide ten recommendations towards better conservation and management of this iconic group. A total of 69 species were assessed as Critically Endangered (8% of all species), 47 as Endangered (6%), 104 as Vulnerable (12%), and 23 as Near Threatened (3%). Slightly over 60% of CPs (521 species) were assessed as Least Concern. At least 89 species are known from only a single location based on current knowledge. Data on threatening processes were available for 790 species, with the most common threatening processes including Agriculture and Aquaculture (impacting 170 species), Natural Systems Modifications (168 species), Climate Change and Severe Weather (158 species), Energy Production and Mining (127 species), Human Intrusions and Disturbance (126 species), and Biological Resource Use (98 species). Almost a quarter of all species were impacted upon by three or more threatening processes. The most significant threats placing species at imminent risk of extinction include the continuing clearing of natural habitat for urban and agricultural development and the illegal collection of individuals from the wild for horticultural trade. The complex and specialised ecological requirements of CPs, together with the multifaceted threats they face, make conservation difficult and repatriation even to restored areas challenging. As the number of vulnerable, endangered and extinct carnivorous plant species continues to grow, despite significant conservation efforts in many regions and greater awareness of their ecological requirements, it is clear that a paradigm shift is required in our approach to the preservation of this unique group of plants in order to achieve long-term conservation successes.
... However since then the Philippines has dominated for discovery of new species. In 2013 alone, twelve new species were published from the Philippines (Cheek & Jebb, 2013a-g), and new species continue to be added every year (Cheek & Jebb, 2014;Cheek et al., 2015;Gronemeyer et al., 2016;Lagunday et al., 2017;Amoroso et al., 2018;Robinson et al., 2019a). The current total of Nepenthes species for the Philippines is 59 (Pelser et al., 2011onwards, accessed May 2020 now far exceeding the totals for other phytogeographic units such as Sumatra and Borneo. ...
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Nepenthes maximoides sp. nov. (Sect. Alatae) is described and assessed as Critically Endangered (Possibly Extinct) from Luzon, Philippines and appears unrecorded in 110 years. The spectacular, large, narrowly funnel-shaped upper pitchers, lids with recurved basal, and filiform apical appendages, unlike any other species in the Philippines, closely resemble those of N. maxima (Sect. Regiae) of Sulawesi-New Guinea, likely due to convergent evolution. Following recent phylogenomic analysis, sect. Alatae is divided into two, Sect. Alatae sensu stricto of Luzon to Sibuyan (including N. maximoides), and Sect. Micramphorae, expanded and recircumscribed to encompass those species of the southern Visayas, and Mindanao. A key is provided to the six species now recognised in the narrowly recircumscribed Sect. Alatae. The number of Nepenthes species recorded from Luzon has increased from two in 2001, to eight in 2020, all but one of which are endemic to that island, and four of which appear to be point endemics.
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Nepenthes maximoides sp. nov. (Sect. Alatae) is described and assessed as Critically Endangered (Possibly Extinct) from Luzon, Philippines and appears unrecorded in 110 years. The spectacular, large, narrowly funnel-shaped upper pitchers, lids with recurved basal and filiform apical appendages, unlike any other species in the Philippines, closely resemble those of N. maxima (Sect. Regiae) of Sulawesi-New Guinea, likely due to convergent evolution. Following recent phylogenomic analysis, sect. Alatae is divided into two, Sect. Alatae sensu stricto of Luzon to Sibuyan (including N. maximoides), and Sect. Micramphorae, expanded and recircumscribed to encompass those species of the southern Visayas, and Mindanao. A key is provided to the six species now recognised in the newly narrowly recircumscribed Sect. Alatae. The number of Nepenthes species recorded from Luzon has increased from two in 2001, to eight in 2020, all but one of which are endemic to that island, and four of which appear to be point endemics.
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Nepenthaceae is one of the largest carnivorous plant families and features ecological and morphological adaptations indicating an impressive adaptive radiation. However, investigation of evolutionary and taxonomic questions is hindered by poor phylogenetic understanding, with previous molecular studies based on limited loci and taxa. We use high-throughput sequencing with a target-capture methodology based on a 353-loci, probe set to recover sequences for 197 samples, representing 151 described or putative Nepenthes species. Phylogenetic analyses were performed using supermatrix and maximum quartet species tree approaches. Our analyses confirm five Western outlier taxa, followed by N. danseri , as successively sister to the remainder of the group. We also find mostly consistent recovery of two major Southeast Asian clades. The first contains common or widespread lowland species plus a Wallacean-New Guinean clade. Within the second clade, sects. Insignes and Tentaculatae are well supported, while geographically defined clades representing Sumatra, Indochina, Peninsular Malaysia, Palawan, Mindanao and Borneo are also consistently recovered. However, we find considerable conflicting signal at the site and locus level, and often unstable backbone relationships. A handful of Bornean taxa are inconsistently placed and require further investigation. We make further suggestions for a modified infra-generic classification of genus Nepenthes .
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A new species, Medinillatheresae Fernando, from ultramafic soils on Dinagat and Mindanao Islands, Philippines is described and illustrated. The species is characterized by its terrestrial erect habit, non-setose nodes, 3-plinerved, lanceolate and coriaceous leaves arranged in whorls, cauline or axillary and pendulous inflorescences, rounded flower buds, 4-merous flowers, and straight anthers. It is compared with other similar species in the Medinillapendula Merr. complex.
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Globally, ultramafic outcrops are renowned for hosting floras with high levels of endemism, including plants with specialised adaptations such as nickel or manganese hyperaccumulation. Soils derived from ultramafic regoliths are generally nutrient-deficient, have major cation imbalances, and have concomitant high concentrations of potentially phytotoxic trace elements, especially nickel. The South and Southeast Asian region has the largest surface occurrences of ultramafic regoliths in the world, but the geoecology of these outcrops is still poorly studied despite severe conservation threats. Due to the paucity of systematic plant collections in many areas and the lack of georeferenced herbarium records and databased information, it is not possible to determine the distribution of species, levels of endemism, and the species most threatened. However, site-specific studies provide insights to the ultramafic geoecology of several locations in South and Southeast Asia. The geoecology of tropical ultramafic regions differs substantially from those in temperate regions in that the vegetation at lower elevations is generally tall forest with relatively low levels of endemism. On ultramafic mountaintops, where the combined forces of edaphic and climatic factors intersect, obligate ultramafic species and hyperendemics often occur. Forest clearing, agricultural development, mining, and climate change-related stressors have contributed to rapid and unprecedented loss of ultramafic-associated habitats in the region. The geoecology of the large ultramafic outcrops of Indonesia’s Sulawesi, Obi and Halmahera, and many other smaller outcrops in South and Southeast Asia, remains largely unexplored, and should be prioritised for study and conservation. Keywords: Adaptations, Conservation, Edaphic endemism, Edaphic flora, Extreme environments, Geobotany, Plant–soil relations, Serpentine vegetation, Ultramafic plants, Metal hyperaccumulators
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With 50 species of the genus Nepenthes L. currently described from the Philippines, it is without doubt that the country, along with the islands of Sumatra (Indonesia) and Borneo (Indonesia, Malaysia, Brunei), should be considered the center of diversity of the genus. In this work, we describe two new species. One species, N. aenigma sp. nov., is from Ilocos Norte province on Luzon Island and has the-for Nepenthes-unusual ecological preference to grow in dense vegetation in deep shade. The other new species is from Mount Hamiguitan in Davao Oriental province on Mindanao Island. With this new entry, Mount Hamiguitan is now home to four endemic species (N. peltata, N. micramphora, N. hamiguitanensis, N. justinae sp. nov.). Furthermore, we provide an emended description of N. ramos based on field data. Nepenthes kurata is synonymized here with N. ramos.
Nepenthes is a genus of carnivorous plants consisting of ~160 species that are distributed in the paleotropics. Molecular systematics has so far not been able to resolve evolutionary relationships of most species because of the limited genetic divergence in previous studies. In the present study, we used a genome-skimming approach to infer phylogenetic relationships on the basis of 81 plastid genes and the highly repetitive rRNA (external transcribed spacer (ETS)–26S) for 39 accessions representing 34 species from eight sections. Maximum-likelihood analysis and Bayesian inference were performed separately for the nuclear and the plastid datasets. Divergence-time estimations were conducted on the basis of a relaxed molecular-clock model, using secondary calibration points. The phylogenetic analyses of the nuclear and plastid datasets yielded well resolved and supported phylogenies. Incongruences between the two datasets were detected, suggesting multiple hybridisation events or incomplete lineage sorting in the deeper and more recent evolutionary history of the genus. The inclusion of several known and suspected hybrids in the phylogenetic analysis provided insights into their parentage. Divergence-time estimations placed the crown diversification of Nepenthes in the early Miocene, c. 20 million years ago. This study showed that genome skimming provides well resolved nuclear and plastid phylogenies that provide valuable insights into the complex evolutionary relationships of Nepenthes.
Nepenthes is a genus of 130-160 species, almost half of which were described after 2001. The recent, rapid increase in species descriptions has been driven by application of a less rigorous species concept by botanists, taxonomic inflation, and discoveries of new taxa during explorations of remote parts of Southeast Asia. Many recently published species descriptions of Nepenthes are based entirely upon qualitative morphological information and are not supported by adequate research. Accordingly, the status of many Nepenthes taxa is contested. Evolution within the genus is not well understood, because nuclear and maternally inherited plastid genomes cannot resolve relationships between many species, particularly those that evolved recently through introgression or reticulate evolution. Improvement in our understanding of the systematics and evolution of Nepenthes requires the adoption of ‘best practice’ collection and preservation methods, and the application of quantitative analytical methods for morphological, genetic, and ecological information.
Nepenthes bellii from Mindanao Island in the Philippines is described as new.