European Journal of Taxonomy 69: 1-23

http://dx.doi.org/10.5852/ejt.2013.69

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2013 • Martin Cheek & Matthew Jebb

Research article

Red rcuinscription of the Nepenthes alata group (Caryophyllales:

Nepenthaceae), in the Philippines, with four new species

Martin CHEEK 1 & Matthew JEBB 2

1 Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, U.K.

Email: [email protected] (corresponding author)

2 National Botanic Garden, Glasnevin, Dublin 9, Ireland

Email: [email protected]

Abstract. An overview of Nepenthes in the Philippines is presented. Four new species, Nepenthes

extincta sp. nov., N. kitanglad sp. nov., N. kurata sp. nov. and N. leyte sp. nov. are described and

illustrated from the Philippines and placed in the Nepenthes alata group. An updated circumscription

and key to the species of the group is provided. Delimitation and comparison with the Regiae group is

given. All four of the newly described species are assessed as threatened using the International Union

for the Conservation of Nature 2012 standard, and one, N. extincta sp. nov. is considered likely to be

already extinct due to open-cast mining. Logging and conversion of forest habitat are thought to be the

main threats to the other three species.

Key words. Conservation, Nepenthes alata group, Mindanao, threatened, ultramafic.

Cheek M. & Jebb M. 2013. Recircumscription of the Nepenthes alata group (Caryophyllales: Nepenthaceae),

in the Philippines, with four new species. European Journal of Taxonomy 69: 1-23. http://dx.doi.org/10.5852/

eit.2013.69

Introduction

This paper forms part of studies towards a World Monograph of Nepenthes L. (Linnaeus 1753)

(Nepenthaceae), building on a Skeletal Revision of Nepenthes (Jebb & Cheek 1997) and the Flora

Malesiana account (Cheek & Jebb 2001).

Nepenthes is the sole genus in its family. The species mainly occur in Malesia, but with ca. 12 outlying

species in Madagascar, Seychelles, Sri Lanka, NE India, Indochina, Solomon Islands, New Caledonia

and Australia. Cheek & Jebb (2001) recorded 87 species of Nepenthes , but since 1 st Jan. 2001, 50

new specific names have been published in the genus (International Plant Names Index continuously

updated) and others have been resurrected from synonymy. The total number of species now accepted in

the genus is estimated as 140 and is set to rise further.

Within the Philippines 12 species were recognised in Cheek & Jebb (2001). Since 1 st Jan. 2001, 16

new names have been published at species-level for the country (International Plant Names Index

continuously updated). These derive mainly from exploration of previously unexplored areas by

Nepenthes enthusiasts (e.g. Heinrich et al. 2009; Cheek 2011) and are also due to the discovery of

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European Journal of Taxonomy 69 : 1-23 ( 2013 )

previously overlooked specimens in herbaria around the world (e.g. Cheek & Jebb 2013a, b, c, d). Some

of the latter, e.g. Nepenthes alzapan Jebb & Cheek (Cheek & Jebb 2013b) are considered possibly

already extinct, since habitat destruction in the Philippines over the last 100 years has been so extensive.

The Philippines is thought to have remained two-thirds forested as recently as 1925 according to Sohmer

& Davis (2007). Myers et al. (2000) estimated that remaining primary vegetation in the Philippines 75

years later amounted to only 3%. Lowland primary forest (occurring below 500 m) has now all but gone

from the Philippines (Sohmer & Davis 2007). Sohmer & Davis (2007) estimate species extinction levels

due to habitat destruction as between 9 and 28% in one representative, mainly forest genus, Psychotria

L. (Linnaeus 1759) (Rubiaceae) which they revised taxonomically for the Philippines. Since habitat

destruction continues in the Philippines, it is a race against time to discover, publish, assess and draw

attention to the conservation needs of species before they become extinct, if they have not already been

lost.

Of the species of Nepenthes known for the Philippines, about half occur in scrub or stunted habitats on

ultramafic substrates (both lowland and submontane) and appear to be restricted to this substrate, while

the remainder occur mainly in submontane forest (500-2000 m) on non-ultramafic, ma inl y volcanic

substrates. Species diversity is highest on Mindanao Island, the southernmost and second-largest island,

followed by Palawan Island.

All known Philippine Nepenthes species are endemic, apart from N. mirabilis Lour. (Druce) (Druce

1916; basionym: Loureiro 1790) which is known from Tawi Tawi Island close to Borneo, and from the

northern coast of Mindanao and its associated islands.

Phytogeographically, the Nepenthes of Palawan have closer affinities with Borneo than with the rest of

the Philippines (Cheek & Jebb 1999).

Most of the endemic species can be assigned to one of three main groups:

(1) The Nepenthes villosa Hoolc.f. (Hooker 1851) group, with petiolate leaves, well-developed lid

columns, and peristomes with blade-like teeth, and lids which lack a basal appendage. These are restricted

to higher altitude ultramafic habitats and are most diverse in Palawan, but extend to NE Borneo (Cheek

& Jebb 1999: 894).

(2) The Insignes or N. ventricosa Blanco (Blanco 1837) group which is most diverse in the Philippines

(but absent from Palawan), extends to New Guinea and to Borneo and is characterised in Cheek & Jebb

(2013b). In the Philippines they occur in both ultramafic and non-ultramafic, ma inl y submontane forest

habitats. Its species lack a petiole, have broadly subcylindrical pitchers with a broad peristome which

is conspicuously toothed on its inner surface, lids which lack appendages, and which have transversely

elliptic nectar glands except along the midline band.

(3) The Nepenthes alata Blanco (Blanco 1837) group, characterised in Cheek & Jebb (2013d), is endemic

to and extends throughout the Philippines except Palawan, is most diverse in Mindanao, and occurs in

the same range of habitats as the Insignes. Its species have winged petioles, lids with basal ridges,

usually producing appendages and upper pitchers usually broadest at the base.

Outside of these three groups, three other species occur, namely N. philippinensis Macfarl. (Macfarlane

1908), N. abalata Jebb & Cheek (Cheek & Jebb 2013a: 153) and N. micramphora V.B.Heinrich,

S.McPherson, Gronem. & V.B.Amoroso (Heinrich et al. 2009). N. philippinensis is endemic to Palawan,

and part of the N. hirsuta Hook.f. (Hooker 1873) group otherwise restricted to Borneo (Cheek & Jebb

1999: 890-891). The unusual N. abalata occurs only in the small low-lying, drier islands between

Palawan and Panay Islands. It appears to be restricted to grassland and is similar and possibly related to

the grassland species of Indo-China. Finally, N. micramphora of the extreme SE of Mindanao fits none

of these groups and has an isolated position.

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CHEEK M. & JEBB M., Nepenthes (Caryophyllales: Nepenthaceae) in the Philippines

Many Philippine species of Nepenthes are only known from a single, sometimes sterile, specimen.

However, usually there is little doubt that they are distinct taxa, since most characters of use in delimiting

species in the genus are found in the pitchers and not in the flowers or fruit. In Nepenthes the large number

of taxonomic characters in the pitcher morphology and micro-morphology, especially in the upper and

intermediate pitchers, parallels the importance of characters present in the flowers and inflorescences of

many other plant groups. In many other plant groups evolution of character diversity in floral characters

appears to be correlated with assisting pollination, often by a single or narrow range of insect species. In

Nepenthes , cross-pollination appears from initial studies to be performed by a broad range of generalist

flies and wasps (Lian 1995; Adam 1998). Perhaps for this reason there is comparatively little species-

level differentiation florally in the family. In contrast, evidence has been steadily accumulating in recent

years that the pitchers of different Nepenthes species are specialised at gathering nutrients from one of a

variety of animal sources. They are not always pitfall trappers of any prey item that happens to fall in, as

had previously been assumed. Nepenthes species studied by Jebb (1991) in New Guinea predominantly

trapped ants, but also a spectrum of other insects, while Nepenthes albomarginata T.Lobb ex Lindley

(Lindley 1849) traps almost exclusively a species of termite (Clarke 1997). Nepenthes hemsleyana

Macfarl. (Macfarlane 1908), the pitchers of which, providing day roosts for a species of bat, benefit

from the resulting faecal deposits produced (Grafe et al. 2011), while a group of Nepenthes species in

montane Borneo are adapted to attract, detain and collect the faeces of a species of tree shrew and a rat

(Greenwood et al. 2011). In each of these cases modification in the morphology and micro-morphology

of the pitchers appears to have been critical to collect nutrients from the target animals.

No doubt in future, morphological and micromorphological character states seen in the pitchers of other

Nepenthes species or species groups, will also be shown to be important to targeting hitherto unsuspected

specialised animal nutrient sources.

In the last three cases, the morphological modifications of the pitcher each appear to target only one or

two species of animal. However, it is possible that future studies will show that some or most species

of Nepenthes trap a wider range of insect species available in their environment. Since the range of e.g.

arthropod species can be expected to vary from one mountain (or other geographic location) to another,

it is possible that speciation in Nepenthes might be driven by modifications of the pitcher caused by

selection for efficiency in trapping, or otherwise targeting, the spectra of animal species present at each.

This might explain the common pattern seen in several Nepenthes groups to have different species, as

recognised by pitcher morphology and micromorphology, on different mountains or islands.

An alternative hypothesis to explain this speciation pattern invokes genetic drift (Stace 1991). In

this hypothesis, the progeny of the few founding individuals at a new location might accumulate

morphological changes over time independent of any selection pressures, for example as postulated

for Podostemaceae Rich, ex Kunth (Kunth 1816) species at the same waterfall by van Steenis (1981),

exemplifying his theory of autonomous evolution. This scenario assumes that animal targeting is not

strongly selected for in Nepenthes. This might be the case if the threshold for the need for nutrients

derived from animals in Nepenthes is very low, or if there is no threshold at all, for example if Nepenthes

can survive without capturing animal nutrients through their pitchers.

Materials and Methods

The data in this study, one of a series, derives from herbarium specimens including their corresponding

field notes, and if available, photographs. The location of the herbarium at which the specimens are

located is indicated by the citation of the international code registered in Index Herbariorum (Thiers

continuously updated).

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European Journal of Taxonomy 69 : 1-23 ( 2013 )

The shape and dimensions of the upper and intermediate pitchers, including those of the body, mouth

and lid of the pitcher, and of the fringed wings if present, are useful characters. The shape and features of

the lower surface of the pitcher lid, and the shape of the pitcher mouth and its peristome are the largest

source of such characters, as are the nectar glands on the lower surface of the lid (Cheek & Jebb 2001).

The nectar gland size, shape, orientation, position, density and the absence or, if present, the extent of

a nectar gland perimeter wall, can be especially useful in delimiting species. In some species nectar

glands are dimorphic, and in a very few, trimorphic. Indumentum is also frequently an important source

of species characters. Here the classification used in Cheek & Jebb (2001: 7-8) is maintained.

Since the mouth and the lid surface are often obscured or not presented in mounted herbarium specimens

of Nepenthes , it is frequently necessary, after prior permission from the herbarium manager of the

specimens concerned, to soak the mouth area and lid, and sometimes entire pitchers, in warm soapy

water so that they can be repositioned for study of the mouth and lower surface of the lid. By this

means it is possible to view the three-dimensional shape of the pitcher, if this is not already known from

photographs or observation of live material.

Measurements are made of stem, leaf and pitcher dimensions using a graduated ruler. The full range of

measurements is given in the description, with extreme dimensions, disconnected with the normal range,

indicated in brackets, e.g. “leaf-blade 12.5-22(-28) cm long”, indicating that a single leaf-blade was

measured as 28 cm long, all others being in the 12.5-22 cm long range. Observations and measurements

of hair dimensions and of peristome and lower lid details are made with a Leica Wild M8 binocular

microscope fitted with an eyepiece graticule graduated in units of 0.025 mm at X40 magnification. The

drawings were prepared with the aid of a camera lucida fitted to the microscope.

The four species described as new in this paper, N. extincta sp. nov., N. kitanglad sp. nov., N. kurata

sp. nov. and N. leyte sp. nov. came to light when examining loans of previously unstudied specimens

from E, BISH and BRIT, which were compared with material from B, BM, BO, F, GH, K, KEP, L,

NDC, NY, PH, S, W, WAG, U, UC, US, on which earlier studies have been published (Jebb & Cheek

1997; Cheek & Jebb 1999, 2001; Cheek 2011; Cheek & Jebb 2013a, b, c, d). The material referred to

matched no other species description, as represented by the both the specimens from the herbaria cited,

and by protologues and other literature regarding the species for which material is only available in

the Philippines and not available for loan. The material best fitted the characters of Nepenthes group

(3) described above, that is the Nepenthes alata group. However, unusual features for this group were

present, which are considered in the discussion at the end of this paper. A key to the species of the group

is provided below.

The rapid increase in the number of new species from Mindanao in recent years is likely to continue as

specimens become available from new areas. Many of the species of the N. alata group on this island

seem narrowly geographically localised, although others, such as N. graciliflora are widespread.

Results

Key to the species of the Nepenthes alata group updated from Cheek & Jebb (2013b)

1. Lower surface of lid, including appendage (if present), densely and evenly covered in uniformly

minute circular nectar glands (0.15-0.2 mm diam.) .2

Lower surface of lid with nectar glands either absent from the appendage (if present) and/or, sparse,

large or dimorphic (larger glands 0.35-0.4 mm diam. or larger).3

2. Stems glabrous to glabrescent; upper pitchers lacking fringed wings in upper part; outer surface

lacking stellate hairs entirely. S LUZON TO MINDANAO. N. graciliflora E lm er (Elmer 1912)

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CHEEK M. & JEBB M., Nepenthes (Caryophyllales: Nepenthaceae) in the Philippines

- Stems persistently pubescent; upper pitchers with fringed wings in upper part; outer pitcher surface

> 50% covered in grey stellate hairs. N LUZON. N. alata Blanco (Blanco 1837)

3. Stem internodes winged from the decurrent petiole bases. MINDANAO, SARANGANIPROV..

. N. saranganiensis Sh.Kurata (Kurata 2003)

- Stem terete or angular, lacking wings.4

4 Upper pitchers funnel-shaped or funneliform-cylindric.5

- Upper pitchers not funnel-shaped, but subcylindric, widest at base, or equally at base and apex .... 6

5. Stems and abaxial surface of midrib moderately densely covered in white appressed hairs 0.5-

1.5 mm long; lid of upper pitchers ovate, longer than broad. Volcanic substrate. MINDANAO, MTS

APO & MATUTUM. N. copelandii Macfarl.(Macfarlane 1908)

- Stems and leaf-blades glabrous; lid of upper pitcher broader than long. Ultramafic substrate.

MINDANAO, MT KIAMO. N. ceciliae Gronem. et al. (Gronemeyer et al. 2012)

6 Petiole appearing flat, at least distally (wings held flat).7

- Petiole appearing cylindrical (wings incurved).12

7. Stems and lower surface of midrib conspicuously densely pubescent; lid appendage well-developed,

hooked. 8

- Stems and inner surface of midrib glabrous or inconspicuously and sparsely pubescent; lid appendage

moderately or well-developed, never hooked.9

8. Upper pitchers widest at base, contracting slightly above into a narrower cylinder with fringed wings.

NEGROS & BILIRAN ISL. N. negros Jebb & Cheek (Cheek & Jebb 2013d)

- Upper pitchers equally wide at base and apex, contracting slightly at the middle, lacking fringed

wings. MINDANAO, SURIGAO PROV.. N. ramos Jebb & Cheek (Cheek & Jebb 2013c)

9. Lid apex with pocket. MINDANAO, S COTABATO. N. tboli Jebb & Cheek sp. ined.

- Lid apex lacking pocket or any appendage. 10

10. Petiole canaliculate proximally (flat distally); blade hairy on upper surface; upper pitchers stout,

length: breadth ratio 2-2.5:1; fringed wings absent. MINDANAO, MT HAMIGUITAN.

. N. hamiguitanensis Gronem. et al. (Gronemeyer et al. 2010)

- Petiole flat proximally; blade glabrous on upper surface; upper pitchers slender, length: breadth ratio

>3:1; fringed wings present below peristome. 11

11. Upper pitcher with lid about half as long as mouth; mouth not concave but flat; column absent; lid

base truncate. MINDANAO, MT MALINDANG. N. kurata Jebb & Cheek sp. nov.

- Upper pitcher with lid about as long as mouth; mouth highly concave; column present; lid base

cordate. MINDANAO, KITANGLAD MTS . N. kitanglad Jebb & Cheek sp. nov.

12. Largest pitchers robust, 18-24 cm long; peristome 7-8 mm wide, curved in section but not cylindric;

inner edge of peristome with small teeth visible, outer edge lobed. MINDANAO.13

- Largest pitchers 12 cm long; peristome 2-3 mm wide, narrowly cylindrical, inner edge lacking

visible teeth (unless dissected), outer edge not lobed. LEYTE ISL. N. leyte Jebb & Cheek sp. nov.

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European Journal of Taxonomy 69 : 1-23 ( 2013 )

13. Largest pitchers without fringed wings; leaf midrib densely, minutely white stellate-hairy; lid base

deeply cordate. MINDANAO, RED MT. N. extincta Jebb & Cheek sp. nov.

- Largest pitchers with fringed wings; leaf midrib with brown black bristle-like hairs 1-1.5 mm long;

lid base truncate. NE MINDANAO. N. mindanaoensis Sh.Kurata (Kurata 2001)

Taxonomic descriptions

Order Caryophyllales Juss. ex Bercht. & J.Presl. sens. lat. (Berchtold & Presl 1820)

Family Nepenthaceae Dumort. (Dumortier 1829)

Genus Nepenthes L. (Linnaeus 1753)

Nepenthes kurata Jebb & Cheek sp. nov.

urn:lsid:ipni.org:names:77134486-1

Fig. 1

Diagnosis

Differs from N. mindanaoensis Sh.Kurata in the petiole wings patent (not involute), the hairs of stem,

midrib and leaf-edge bushy, 0.1 mm long, not bristle-like 1-1.5 mm long; the lid about half as long as

the mouth, lid base rounded or truncate (not about as long as the mouth, base cordate).

Etymology

Named as a noun in apposition for Shigeo Kurata, whose book on the Nepenthes of Mount Kinabalu

(Kurata 1976) inspired interest in the genus among its many readers, and whose descriptions of Nepenthes

are models of detail, precision and clarity.

Type

PHILIPPINES. Mindanao, “Prov. of Misamis, Mount Malindang”. May 1906, Mearns & Hutchinson in

Forest Bureau 4632 (holotype K!; isotype PH!). Fig 1.

Synonym

Nepenthes alata Blanco var. ecristata Macfarlane, Nepenthaceae. In: Engler A. (ed.) Das Pflanzenreich

Heft 36, 4, 3: 72 (1908). - Type: lectotype, designated here: Philippines, Mindanao, “Prov. of Misamis,

Mount Malindang”, May 1906, Mearns & Hutchinson in Forest Bureau 4632 (lecto-: K!; isolecto-:

PH!).

Description

Terrestrial shrub-cl imber. height unknown. Climbing stems terete to slightly angular, 4-6 mm diam.;

internodes 30-50 mm long; axillary buds not evident; indumentum inconspicuous, persistent to the

fifth internode from the apex, hairs translucent brown, simple or 2-3-armed from the base, hairs

straight, variously angled from the horizontal, ca. 0.1 mm long, covering ca. 5% of the surface except

the axils (100% coverage) surface brown-black, matt. Leaves of rosette shoots thinly coriaceous,

blade narrowly elliptic, 8-9 x 2-2.5 cm; apex and base acute; longitudinal nerves 1-2 pairs, within 2

mm of the margin, moderately conspicuous on both surfaces; pennate nerves at 90° from the midrib,

numerous and moderately conspicuous; upper surface drying glossy pale brown, lower surface matt,

mid-brown. Leaves of climbing stems as the rosette leaves, but blades suboblong or oblong-lanceolate

10-12.5 x 3.2-3.8 cm; apex obtuse or acute; base obtuse; lower surface with sessile red glands ca.

0.5 mm diam.; midrib 40-60% covered in patent, brown, simple or basally bifurcate-trifurcate hairs

0.1—0.3(—0.5) mm long; margin fringed, in young leaves, with hairs 0.25 mm long, pale-brown,

1 ^1-armed from the base. Petiole winged-canaliculate, 4-5 x 0.7 cm, wings patent; base clasping the

stem for A to A its circumference, sometimes decurrent as an obtuse ridge to the node below. Lower

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CHEEK M. & JEBB M., Nepenthes (Caryophyllales: Nepenthaceae) in the Philippines

Fig. 1. Nepenthes kurata sp. nov. A. Habit, climbing stem with upper pitcher. B. Indumentum of midrib,

lower surface of leaf-blade. C. Indumentum of tendril. D. Indumentum of outer pitcher surface. E.

Lid, lower surface, showing nectar gland distribution on right (upper pitcher). F-G. Profiles of basal

lid ridges, without appendage (F), with appendage wealdy developed (G), and moderately developed

(H). I Spur of intermediate pitcher. J. Longitudinally elliptic nectar glands of lid midline. K. Orbicular

nectar glands of lid, outside the midline. L. Peristome from above, short teeth and holes discernible. M.

Peristome viewed from inside pitcher. N. Peristome transverse section, outer surface to right. A-E, J &

K from Gaerlan et al. PPI109IP, F-I, L-N from Mearns & Hutchinson 4632. Scale bars: single = 1

mm; graduated single = 2 mm; double = 1 cm; graduated double = 5 cm. All drawn by Andrew Brown.

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European Journal of Taxonomy 69 : 1-23 ( 2013 )

pitchers unknown. Intermediate pitchers (tendrils uncoiled: Mearns & Hutchinson 4632 ) 12.5-17.2 cm

long, ellipsoid in the basal third to half, 4-5.7 cm wide, constricted, more or less abruptly, 5-7.5 cm

from the base into the subcylindrical upper part, 2.1-3 cm diam. dilating slightly towards the apex

3-4 cm diam.; outer surface strongly reticulated with raised nerves when dry, 2-5% covered in hairs

of two types (Fig. ID), (1) large erect hairs 0.3-0.75 mm long, with a single, major, curved arm, and

1-2 much smaller erect arms, and (2) minute, 3-6-armed stellate hairs 0.05-0.1 mm diam., which are

more frequent, (ca. 4 per mm 2 ); surface covered throughout (6-10 per mm 2 ) with sessile, depressed-

globose glands 0.1-0.2 mm diam.; fringed wings reduced to ridges except in the ca. 25 mm below the

peristome, widening to 3 mm broad, with fringed elements 2.5 mm long, 2-5 mm apart; mouth oblique,

suborbicular, ovate, 3—4.8 x 2.7—4.5 cm; apex with a column 9-10 mm long; peristome rounded to

slightly flattened, 2-2.5 mm wide, more or less even in width, ribs 0.25-0.5 mm apart, conspicuous,

about 0.1 mm high, outer edge lacking lobes, inner edge with very short teeth and conspicuous holes,

teeth <0.1 mm long. Ljfl much smaller than the mouth, ovate, or broadly ovate, 25-35 x 25-30 mm,

apex rounded to obtuse, base rounded to truncate; lower surface with a low basal ridge ca. 1 mm high,

7-10 mm long, either lacking a protruding appendage entirely (Fig. IF) or with a modestly developed

appendage 1-2 mm high (Fig. 1H); nectar glands only slightly dimorphic, (1) midline nectar glands

sparse, longitudinally elliptic, 0.5-0.7 x 0.1-0.25 mm, with a thin marginal rim (Fig. 1J), (2) outside the

midline nectar glands circular (Fig. IK), sparse, < 1 per mm 2 , only 35-50 on each side of the midline,

the largest scattered in the distal half, 0.5 mm diam., grading down to those of the marginal equatorial

areas ca. 0.25 mm diam., and those at the attachment point with the peristome and the basal ridge and

appendage, 0.15 mm diam.; sessile depressed-globose minute red glands 0.1-0.2 mm diam. are scattered

over the surface at a density of 3-8 glands per mm 2 ; minute inconspicuous stellate hairs ca. 0.075 mm

diam. occur in an uneven, 0.5-1 mm wide band, near the margin widening to 1.5 mm wide at the lid

apex. Spur unbranched, curving downwards, stout at base and tapering to a slender apex, ca. 5 mm long,

with scattered long, subpatent hairs 0.3-0.7 mm long (Fig. II). Upper pitchers (tendril coiled, Gaerlan

et al. in PPI 10914 ) resembling the intermediate pitchers, but fringed wings 1-2 mm wide, fringed

elements 2.5 mm long, (2-)4-5 mm apart, dilating to 4.5 cm below the mouth; pitcher green, peristome

maroon. Lid broadly ovate to suborbicular 32 x 35 mm, lower surface with a basal ridge 9-10 mm long,

ca. 2 mm high, bearing a central, symmetrical, protruding appendage 2x3 mm; nectar glands denser, ca.

110 on each side of the midline. Male and female inflorescences and infructescences unknown.

Additional material

PHILIPPINES. Mindanao, Prov. Misamis Occidental, S.E. slopes of Mt. Malindang, Lake Duminagat,

May 1993, Gaerlan, Sagcal & Romero in PPI 10911 (BRIT!).

Distribution, habitat & phenology

Philippines, Mindanao; evergreen forest, volcanic substrates. Elevation: ca. 1400 m.

Conservation status

Nepenthes kurata sp. nov. is here assessed as Critically Endangered under Criterion D of IUCN (2012)

since currently only two individuals, probably at a single location (as currently defined by IUCN)

are known. This site, the ca. 6 ha crater Lake Duminagat, is within the ca. 50,000 ha Mt Malindang

Range Natural Park of which at least 20,000 ha has been cleared for cultivation purposes, but which

is a tentative World Heritage Site ( http://whc.unesco.org/en/tentativelists/5029/ . downloaded 16 July

2013). In 2012 the Park was designated as an ASEAN Heritage Park ( http://news.pia.gov.ph/index.

php?article= 1451343449808 . downloaded 16 July 2013). It is to be hoped that further investigation will

discover additional individuals and locations for this species, decreasing its threat status, and increasing

the likelihood that it can be protected.

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CHEEK M. & JEBB M., Nepenthes (Caryophyllales: Nepenthaceae) in the Philippines

Since the terrain of Mt Malindang is reported as being rugged, with much forest surviving, there is every

reason to hope that the species survives there, unlike Nepenthes robcantleyi Cheek (Cheek 2011) also

from Mindanao, which is already suspected to be extinct in the wild due to the almost total clearance of

forest habitat at the single known wild location due to logging (Cheek 2011).

Remarks

The first Nepenthes taxa described from Mindanao, both of the N. alata group, (Cheek & Jebb 2013d),

were N. alata var. ecristata Macfarl. (Macfarlane 1908), based on Mearns & Hutchinson 4632 from Mt

Malindang, and TV. copelandii Macfarl. (Macfarlane 1908) fromMtApo.

The first of these we here elevate to species level as N. kurata Jebb & Cheek sp. nov. Previously we

had considered this taxon to be synonymous with N. mindanaoensis Sh.Kurata (Kurata 2001) (Cheek

& Jebb 2013d). The two taxa do have similarities in the overall shape of the upper pitchers, the weakly

to moderately developed basal lid appendage and the sparse nectar glands of the lower lid surface.

However they can be distinguished using the characters in Table 1. The number and extent of these

features merit elevation from varietal to specific-level recognition in our opinion. Although the type

specimen has rosette stems and intermediate pitchers only, a second specimen, with climbing stems and

upper pitchers, Gaerlan et al. in PPI10911 came to light recently. It is from the type locality and matches

the type in essential details. Nepenthes kurata sp. nov. has the spot character within the Nepenthes alata

group of a small, more or less orbicular lid, only about half the length of the pitcher mouth.

Macfarlane (1908) characterised his N. alata var. ecristata by the lid appendage being either reduced

or absent; the nectar glands being few, medium to large in size, and irregularly dispersed. Of the single

specimen cited ( Mearns & Hutchinson 4632), only two sheets (PH and K) have been found, both

annotated in Macfarlane’s hand, each with two intermediate pitchers. Although all four pitchers share a

basal ridge (Fig. 1F-H), only one of the four has an appendage, and that is only moderately developed as

a convex emergence from the basal ridge (Fig. 1H). However a recent collection ( Gaerlan et al. in PPI

10977 ) with upper pitchers, does show a developed appendage (Fig. IE), suggesting the epithet ecristata

“lacking a crest” is inappropriate. In any case, the Code demands priority only at one rank, so there is

no requirement to adopt the varietal epithet at specific level, for which reason Macfarlane’s taxon is re¬

named as N. kurata sp. nov.

The upper pitchers also differ from the intermediate pitchers in the greater density of the nectar glands

on the lower surface of the lid. However the shape, distribution and size of the nectar glands remain

s im ilar

This is the only known species of Nepenthes from Mt Malindang at this time, and it is therefore the most

westerly known species of the genus in Mindanao. Nepenthes kurata sp. nov. is still incompletely known,

full details on its ecology, altitudinal range, population density, inflorescences and infructescences, and

ethnobotany remain to be discovered.

The type specimens were collected by Major E.A. Mearns and W.J. Hutchinson in 1906 on the first

recorded ascent of Mt Malindang, a volcanic mountain in the NW of Mindanao. Both sheets are

annotated in the hand of Macfarlane as “A. alata var. ecristata Macfarlane”, and either could be selected

as lectotype of that name. The K sheet is accordingly selected.

9

European Journal of Taxonomy 69 : 1-23 ( 2013 )

Table 1. Diagnostic characters separating N. kurata sp. nov. and N. mindanaoensis.

Nepenthes kurata sp. nov.

Nepenthes mindanaoensis

Geography

Mt Malindang, NW Mindanao

Surigao Prov. NE Mindanao

Substrate

Volcanic

Ultramafic

Habitat m a.s.l.

Submontane forest; ca. 1400 m

Open scrub; 200-300 m

Indumentum (stem, leaf

midrib and margin

Translucent-brown, simple or

bushy, 0.1 mm long

Dark red-brown, bristle-lilce hairs

1-1.5 mm long

Lower surface leaf-blade

Hairs absent (except midrib)

Hairy

Petiole wings

Patent

Involute (petiole appearing

cylindrical)

Longitudinal nerves of leaf-

blade

Arising along length of midrib

Arising from petiole

Pitcher lid base

Truncate or rounded

Deeply cordate

Mouth:lid (length ratio)

ca. 2:1

0.9-1.2:1

Midline nectar glands of

lower lid surface

Longitudinally elliptic (versus

circular outside midline)

Circular, not differentiated from

those of the rest of the lid

Nepenthes kitanglad Jebb & Cheek sp. nov.

urn:lsid:ipni.org:names:77134487-1

Fig. 2

Diagnosis

Differs from N. saranganiensis Sh.Kurata in having angled, (not winged) stems, lower and upper pitchers

strongly dimorphic, (not sub dimorphic); in being a climbing epiphyte of forest (not a terrestrial shrub of

open areas) and in having a strongly concave pitcher mouth with a long neck (not with the pitcher mouth

flat or only slightly concave, lacking a neck).

Etymology

Named, as a noun in apposition, for Mt Kitanglad, the type and only known locality of the species.

Type

PHILIPPINES. Mindanao, Bukidnon Province, Intavas, Impasug-ong, Mt Kitanglad, 18 Jul. 1991,

Gaerlan, Sagcal & Fernando in PPI3274 (holotype BISH!; isotype BRIT!).

Description

Epiphytic climber , probably 1 m tall or more. Short stems terete, 4-5 mm diam., internodes ca. 6 cm

long, axillary buds not evident; surface glossy, appearing glabrous but with extremely sparse brown,

simple hairs ca. 0.5 mm long, glabrescent. Climbing stems strongly 4-angular, 7-9 mm diam., internodes

11-12 cm long; indumentum as short stems. Leaves of short, and of climbing stems more or less identical,

thickly papery; blade oblong-elliptic, 15.5-33 x 4.3-7 cm; apex acute, not peltate, tendril arising

abruptly; base cuneate, decurrent to petiole; longitudinal nerves 3-4 pairs, conspicuous in the marginal

half on the upper surface; pennate nerves arising at about 45° from the midrib, irregular, reticulate,

branching in the marginal half; drying brown-black above, matt mid-brown below, appearing glabrous

apart from margin but with indumentum as stem, densest on midrib but soon glabrescent; lower surface

with sessile depressed-globose red-black glands ca. 0.05 mm diam.; margin densely fringed with soft

fine orange-brown patent simple or bifurcate hairs 1 mm long. Petiole evenly winged along its length,

4-5 x 0.4-0.9 cm, wings patent; at base clasping the stem for 2 /s to 3 /4 of its circumference, decurrent

10

CHEEK M. & JEBB M., Nepenthes (Caryophyllales: Nepenthaceae) in the Philippines

<SAPfe

Fig. 2. Nepenthes kitanglad sp. nov. A. Habit, climbing stem with upper pitcher (from herbarium

specimen). B. Upper pitcher (from live plant). C. Lower pitcher. D. Indumentum from leaf margin. E.

Indumentum from tendril. F. Indumentum from stem (extremely sparse hairs). G. Indumentum of outer

pitcher surface, plan view. H. Ibid., profile view. I. Lower surface of lid (upper pitcher). J. Detail of

glands from lower surface of lid. K. Spur of upper pitcher. L. Peristome, upper pitcher, view from above.

M. Peristome, view from inside of pitcher, showing minute teeth. N. Peristome, transverse section (outer

surface on right). A, D-N from Gaerlan et al. in PPI3274 (BISH); B from McPherson (2009: Fig 417);

C from Gaerlan et al. in PPI 3274 (BRIT). Scale bars: double = 1 cm; double graduated = 5 cm; single

= 1 mm; single graduated = 2 mm All drawn by Andrew Brown.

11

European Journal of Taxonomy 69 : 1-23 ( 2013 )

diagonally as a narrow wing, in short stems 7 mm long, in climbing stems 18 mm long, and continuing

as a ridge to the node below. Lower pitchers narrowly ovoid-cylindric, 12.5 cm tall, 5 cm broad, widest

in the basal half, narrowing steadily to ca. 3 cm wide below the peristome; outer surface 10-25%

covered in pale brown hairs of two types, (1) bushy brown hairs 0.1-0.25 mm long and wide, with 4-8

arms ascending from a short central axis, 7—12 per mm 2 , (2) long brown straight erect hairs 1.5-1.75

(-2.5) mm long, with 2-4 short branches ascending from along the length of the main axis, sparse;

fringed wings, 2-4 mm wide, running 3-4 cm from peristome towards base of pitcher, then d imini shed

to slender ridges, wings extended over the peristome by two foliose flaps 3-4 x 3-4 mm, fringed

elements 4-5 mm long, 2.5 mm apart (1.5 mm apart on foliose flaps); mouth ovate-lanceolate, highly

oblique, concave, ca. 4.1 x 2.8 cm; column developed, tapering towards lid ca. 9 mm long, 2.5 mm wide

at midpoint; peristome subcylindric, 1 mm wide at front of pitcher to 3 mm wide at sides, ridges ca.

2.5 per mm, ridges 0.1 mm high, inner edge lacking conspicuous teeth or holes, outer edge not lobed. Lid

narrowly ovate to rhombic 3.5 x 2.3 cm, apex rounded, base rounded to truncate, lower surface lacking

a basal appendage, but with a low basal ridge 10 mm long, 0.5-1 mm high, extending from the junction

with the peristome; nectar glands small and sparse, 6-8 on each side of the midline which mainly lacks

glands, absent from basal ridge, nectar glands monomorphic, slightly perithecoid, orbicular or slightly

elliptic, 0.25(-0.35) mm long, mixed with denser sessile depressed-globose, red-black glands, 0.05-

0.1 mm diam., 8 per mm 2 ; marginal 2-3 mm of lower surface with minute stellate hairs densest near

margin; upper surface with same indumentum as outer pitcher surface, but long hairs rarely seen. Spur

not seen. Upper pitchers (tendril coiled) ovoid-cylindric, green, slightly maroon above, 21.5 x 6.5 cm,

widest in the ovoid basal third, narrowing to ca. 5 cm wide in the cylindrical upper part; outer surface

with same indumentum as lower pitcher; fringed wings reduced to ridges apart from two foliose flaps

immediately below peristome, point of attachment 3M mm long, angular-elliptic, 9^6 mm, bearing

fringes 2-7 mm long; mouth ovate-lanceolate 7x4 cm, oblique, concave, the frontal part straight;

column ca. 1.5 x 0.8 cm; peristome rounded-flattened, 1.75-5.5 mm wide, widest at sides, ca. 1.75 ridges

per mm, ridges 0.1 mm high, inner edge lacking conspicuous teeth or holes, outer edge not lobed. Lid

ovate-triangular, 5 x 3.8—4.2 cm, apex rounded, base truncate; lower surface with a weakly developed,

convex basal appendage 1.5 mm high, arising from a low basal ridge 7 mm long; nectar glands ca.

16 on each side of the midline, sparsely scattered, more or less absent from midline, but present at

appendage, nectar glands orbicular or slightly elliptic, slightly or strongly perithecoid, 0.25-0.5 x

0.25-0.45(-0.75) mm; sessile glands 0.05-0.1 mm diam., 8-20 per mm 2 ; upper surface of lid with

indumentum as outer surface of pitcher. Spur inserted 2 mm below junction of lid and pitcher, pointing

downwards, terete, 17 x 0.9-1 mm, dilating to the 1.8 mm wide rhombic-acute apex, indumentum

moderately dense of long patent simple hairs as on the pitcher outer surface. Male and female

inflorescences unknown.

Distribution and habitat

Philippines, Mindanao, Bukidnon Province, known only from Mt Kitanglad; epiphytic in mossy forest,

geology volcanic, elevation 1800-2100 m.

Conservation

Here N. kitanglad sp. nov. is assessed as Critically Endangered since it is known from only a single

location, Mt Kitanglad, on an island which has seen extensive forest clearance for logging and

agricultural expansion in recent years (McPherson 2009: 759). For these reasons one species, N.

robcantleyi Cheek is already suspected to be extinct in the wild (Cheek 2011). Nepenthes kitanglad sp.

nov. is not a spectacular or especially bizarre species so is unlikely to come under pressure of collection

for the horticultural trade which has brought several species of the genus close to extinction.

12

CHEEK M. & JEBB M., Nepenthes (Caryophyllales: Nepenthaceae) in the Philippines

Remarks

McPherson (2009: 755-759; figs 417 & 418) depicts from volcanic Mt Kitanglad inN-Central Mindanao

a plant under the name of N. saranganiensis Sh.Kurata (Kurata 2003: 41). Yet, the Kitanglad plants he

depicts differ from N. saranganiensis as depicted in its protologue in habit, habitat and in morphology. In

2013 sheets of Gaerlan etal. inPPI3274 (BISH, BRIT) became available from Kitanglad. These matched

McPherson’s (2009) depiction, enabling a detailed comparison to be made with N. saranganiensis. The

conclusion is that the Kitanglad material represents a different species from N. saranganiensis and is

here described as N. kitanglad sp. nov. Differences between the two taxa are given in Table 2.

N. kitanglad sp. nov. is unusual in the N. alata species group (Cheek & Jebb 2013d) in the strongly

concave mouth of the upper pitchers, in which the base of the lid is held over the mouth. It is also

unusual in that the rear of the peristome narrows to a neck, forming a moderately well-defined column

for the lid. Within the N. alata group, these two features are otherwise currently known only in N.

hamiguitanensis Gronem., Wistuba, V.B.Heinrich, S.McPherson, Mey & V.B.Amoroso (Gronemeyer

et al. 2010), but that species is restricted to ultramafic Mt Hamiguitan in SE Mindanao and differs

greatly in the shape of the upper pitchers which are stout, widest at the midpoint, with a funneliform

lower half narrowing to a more slender, cylindrical upper half. It is possible that N. hamiguitanensis ,

N. kitanglad sp. nov. and N. saranganiensis are related since all have angled stems (or in the case of

the last species, winged), a feature otherwise unknown in the otherwise terete-stemmed N. alata group.

Nepenthes kitanglad sp. nov. is unique in the N. alata group in its lid posture, as seen from photographs

and herbarium specimens. The lid is held at ca. 45° below the horizontal, largely concealing the mouth.

In other species it is usually elevated above the horizontal, sometimes by ca. 45° (TV. saranganiensis)

or as much as 90° or more (TV. graciliflora Elmer). So far, N. kitanglad sp. nov. is the only species of

Nepenthes recorded from Mt Kitanglad.

Table 2. Diagnostic characters separating N. saranganiensis and N. kitanglad sp. nov.

TV. saranganiensis

TV. kitanglad sp. nov.

Habit

Terrestrial shrub

Climbing epiphyte

Habitat

Open, shrubby areas

Forest

Altitudinal range

700-800 m

1800-2100m

Climbing stems

Terete, winged, wings 1.5-2 mm

wide

Angled, wings absent

Lower & upper pitchers

dimorphic?

Monomorphic-subdimorphic

Strongly dimorphic

Mouth

Ovate, flat to slightly concave

Lanceolate; strongly concave

Column

Absent

Present, 0.9 cm long

Lid posture

Elevated above horizontal by

ca. 45°

Declinate below horizontal by

ca. 45°

Lid nectar gland

Midline dense with large glands

Midline ± lacking glands

Lid basal ridge and

appendage

Convex appendage present

Ridge only, appendage absent

Upper pitcher shape

Equally wide at base and apex,

waisted at midpoint

Basal third widest, ovoid, tapering

into the narrow, cylindric upper

part

13

European Journal of Taxonomy 69 : 1-23 ( 2013 )

Nepenthes extincta Jebb & Cheek sp. nov.

urn:lsid:ipni.org:names:77134488-1

Fig. 3

Diagnosis

Differs from N. mindanaoensis Sh.Kurata in the pitchers lacking fringed wings (versus with fringed

wings), the lid base truncate (not cordate), the indumentum of the midrib of dense minute grey-white

stellate hairs, (not of sparse black bristle-lilce hairs).

Etymology

Nepenthes extincta sp. nov. is named to signify that this species may already be extinct globally.

Type

PHILIPPINES. Mindanao Island “Red Hills (= 400 m alt.), SE of Claver, near the northeastern coast of

the Mindanao Island. Boundary of the Surigao del Sur and Surigao del Norte” 8 Aug. 1978, Fumihiro

Konta 12365 (holotype BISH!).

Description

Terrestrial shrub, probably about 1 m tall. Leaves elliptic to elliptic-lanceolate, 13-17 x 5.5-8 cm,

thickly leathery, glossy above, matt mid brown below; apex obtuse to acute, not peltate; base rounded to

obtuse, not decurrent; longitudinal nerves arising from base of blade, where 5-6 pairs arise on each side

of the midrib, at blade midpoint 4-5 pairs occur in the outer third of the blade; pennate nerves arising at

ca. 45° from the midrib, irregularly branching, ends traversing the inner longitudinal nerves; all nerves

most conspicuous on upper surface; midrib deeply depressed on upper surface, lacking hairs, highly

exserted on lower surface and densely (80-90% cover) grey-white stellate hairy, the hairs gathering

dirt, hairs sessile, arms 5-8, 0.25-0.5 mm diam., fine, acute, appressed to surface; lower surface of

blade sparsely hairy, densest towards midrib, ca. 15% cover, decreasing at margin to 5-10%, cover,

hairs mainly stellate, as midrib, mixed with sparser erect, bristle-like hairs 1-2 mm long, of several

types (1) with short branches arising along the length of the main axis; (2) with 2-6 ± equal erect

arms; (3) with a single long erect bristle arising from a stellate hair, the “dagger-hair” of Kurata (2003)

more rarefy (4) hairs with 2-3 erect, equal arms from the base; depressed-globose sessile red-black

glands 0.03 mm diam., raised, dense, conspicuous; upper surface of blade with stellate hairs, as lower

surface, scattered along the margins of the midrib. Petiole 4.5 x 0.5-0.7cm, appearing cylindric due to

the two involute wings, indumentum of appressed, stellate, fine 5-8-armed hairs, ca. 20% cover. Lower

and upper pitchers unknown, possibly not produced. Intermediate pitchers (tendrils not coiled, fringed

wings absent) ovoid-cylindric, 18-24 x 5.9-8.2 cm, widest in the basal half, narrowing gradually to

the cylindric upper half (4.8-5.5 cm wide), not constricted or waisted; outer surface 10-20% covered

in minute, white, 2—4-armed, bushy-stellate hairs 0.12-0.15 mm wide and high, the arms stout and

raised, 10-15 hairs per mm 2 , mixed with sparser black depressed-globose sessile glands 0.07 mm

diam., 4-5 per mm 2 , long simple and bristle-like hairs absent; fringed wings absent, reduced to ridges;

mouth ovate 7-8 x 4.5-5.5 cm, oblique, concave, column weakly defined ca. 1.5 x 0.7 cm; peristome

subcylindric, 7-8 mm wide, widest at sides, outer edge lobed, lobes 1-3 per side, 10-12 mm wide, inner

side inconspicuously toothed, teeth 0.25 mm long; ridges 4 per mm, 0.1 mm high. Lid ovate 5.2-6.5 x

4-5.2 cm, apex rounded, base truncate, lower surface with a basal ridge ca. 8 mm long, 1-2 mm high,

bearing a pronounced straight convex appendage 4 x 2.5-5 mm; nectar glands of two distinct size

classes (1) smaller, elliptic or orbicular, frequent, bordered glands 0.3-0.6(-0.7) x 0.25 mm, the border

ca. 0.1 mm wide, glossy pale brown, dense, (l(-2) per mm 2 ) along the midline these are longitudinally

elliptic, elsewhere with their short axis orientated towards the base of the lid; the appendage completely

covered in smaller type nectar glands; (2) larger glands elliptic to orbicular, (1-) 1.25-2 x 1.25 mm, the

14

CHEEK M. & JEBB M., Nepenthes (Caryophyllales: Nepenthaceae) in the Philippines

QAPb.

Fig. 3. Nepenthes extincta sp. nov. A. Habit, showing intermediate pitcher. B. Indumentum, lower

surface of blade (midrib on left). C. Branched bristle hairs (detail from lower surface of blade). D.

Stellate hairs and sessile glands (detail from lower surface of blade). E. Branched erect hairs (detail

from lower surface of blade). F. Stellate hairs, with appressed arms, leaf-blade lower surface, near

midrib. G. Upper part of intermediate pitcher, lid posture as in life. H. Lower surface of upper pitcher

lid. I. Basal lid appendage and spur. J. Basal lid appendage and ridge, side view. K-M. Details of nectar

glands, lower lid surface. N. Indumentum, outer surface of pitcher. O. Peristome, view from inside

pitcher showing minute teeth and holes at edge. P. Peristome, view from above. Q. Transverse section

of peristome (outer surface on right). A-F & H-Q from Konta 12365 , G from sketch by M. Cheek. Scale

bars: double, graduated = 5 cm; single = 1 mm; single graduated = 2 or 3 mm as indicated. All drawn

by Andrew Brown.

15

European Journal of Taxonomy 69 : 1-23 ( 2013 )

lumina often invaginated by a projection of the border, border 0.2 mm thick, 4-15 on each side of the lid,

scattered around the margin and towards the apex of the midline; sessile, depressed-globose, red glands,

0.05 mm in diam., 1-2 per mm 2 ; marginal 0.5-1 mm of lower surface with minute branched hairs 0.1 x

0.1 mm; upper surface with indumentum as outer pitcher. Spur inserted 5-6 mm below junction of lid

with peristome, cylindric 8-14 x 1-1.2 mm, apex shortly bifid, surface covered in minute appressed,

matted, white-grey stellate hairs. Male and female inflorescences and inffuctescence unknown.

Distribution & habitat

Philippines, Mindanao, Surigao del Sur. Open scrub habitats on ultramafic substrate with N. merrilliana

Macfarl. (Macfarlane 1911) and N. graciliflora Elmer (Elmer 1912). Elevation: ca. 400 m.

Conservation

Nepenthes exincta sp. nov. is here assessed as Critically Endangered under Criterion D of IUCN (2012)

since only a single individual has ever been recorded (the type specimen collected in 1978). The locality

data given corresponds with the large area of ultramafic known as ‘Red Mountain’ SE of Claver in NE

Mindanao. In fact, Red Mountain is a series of low red hills. The NE Mindanao, probably due to its

large areas of ultramafic substrate, supports several narrowly endemic and often spectacular Nepenthes

species, several of which are known from single locations. For this reason, it has been intensively visited

in recent decades by devotees of the genus. Despite this, no additional collections or observations of this

taxon have come to light in the last 25 years and it is possible that it is restricted to the Red Mountain,

and is now extinct, since this happens to be the largest nickel mining site in the world’s second largest

nickel producing country (Gallares 2013). The Foundation for the Philippine Environment (Pacudan

2013) recently reported on the environmental damage due to extensive and massive nickel mining “as

far as your eyes can see.”, “The scene of endless open pit mining at the red mountain made our heart

bleed.” (Pacudan 2013). The biggest mining companies operating at the Red Mountain are Taganito

Mining Corporation, Platinum Group Metals Corporation (PGMC), Taganito HPAL Nickel Corporation,

Adnama Mining Resources Inc. (AMRI) and Zhen Shou Mining (Almeda 2012). It is much to be hoped

that this species is refound, and not proved to be extinct, and if found, that it can be protected and

monitored.

Remarks

Nepenthes extincta sp. nov. is most likely to be confused with N. mindanaoensis and they may share a

common origin. Both species occur on open ultramafic substrates in NE Mindanao, both have robust,

ovoid-cylindric pitchers arising from thickly leathery blades in which the longitudinal nerves arise from

the base of the blade. In both species the petiole has involute wings, so appear cylindric, and the blade is

not decurrent to the petiole - unusual features in the N. alata group. The two can be distinguished using

the characters in Table 3. The two species are not sympatric so far as is known.

Fumihiro Konta, collector of the only known material of this species, has not been traced by us. An

internet search shows that he has published on the plants of S China and Thailand, as well as Japan,

covering specialisms such as Asclepiadaceae, Ferns and Vegetation mapping. “A list of the Ferns and

Flowering Plants of Mt Fuji, 1984” is his most referenced work. Since “Environmental Impact Studies”

are listed among his interests, it may have been in that context that he collected the type specimen

recorded here.

16

CHEEK M. & JEBB M., Nepenthes (Caryophyllales: Nepenthaceae) in the Philippines

Table 3. Diagnostic characters separating N. extincta sp. nov. from N. Mindanaoensis.

Nepenthes extincta sp. nov.

Nepenthes mindanaoensis

Indumentum of leaf-blade

(lower surface of midrib)

Densely covered in grey-white

stellate hairs 0.25-0.5 mm diam.,

arms 5-8, fine, appressed to

surface

Sparsely covered in brown or

black, erect bristle-like hairs

1—1.5 mm long

Intermediate/Upper Pitchers

shape

Ovoid basal half gradually

tapering into cylindric upper half

Ovoid basal third abruptly

narrowing to cylindric upper two-

thirds

Fringed wings

Absent from intermediate pitchers

Present on lower, intermediate

and upper pitchers

Lid base

± truncate

Markedly cordate

Basal appendage of lid

Strongly developed

Absent

Nepenthes leyte Jebb & Cheek, sp. nov.

urn:lsid:ipni.org:names:77134489-1

Fig. 4

Diagnosis

Differs from N. alata B lanco in having, except in the hairy axils, glabrous stems, (not densely white

hairy); upper pitchers lacking fringed wings (versus with fringed wings); nectar glands on lower surface

of lid dimorphic, concentrated around margin and appendage (not monomorphic, uniformly dense and

distributed).

Etymology

The specific epithet “leyte” is here used as a noun in apposition, to commemorate the island of that

name, to which the species appears unique.

Type

PHILIPPINES. Eastern Visayas, Leyte, Mt Lobi, near Dagami, 7 Nov. 1999, Argent, Mendum, Fuentes,

R, Belonias, B.S. 99214 (holotype K!; isotypes E!, PNH).

Description

Terrestrial climber , height 2 m (probably), drying brown. Climbing stems subterete, 4-6 mm diam.,

with a slight ridge below the leaf bases; the axil with a shallow groove containing a spilce-like bud

1-2 mm long, inserted 5-6 mm above the axil; internodes 3-7 cm long; surface with scattered red-

black, depressed-globose, sessile raised glands 0.05-0.08 mm diam.; hairs absent, except in the axillary

grooves which have white, moderately dense, basally branched hairs with arms erect, ca. 1 mm long.

Rosette stems and leaves unknown. Leaves of climbing stems spirally inserted, thinly leathery; blade

narrowly oblong-elliptic, 13.5-16 x 2.5-3.8 cm; apex acute, not peltate; base cuneate, abruptly decurrent

to the petiole; longitudinal nerves 1 pair, moderately close to the margin, inconspicuous; pennate nerves

numerous, conspicuously raised on both surfaces, more or less patent, irregular; both surfaces drying

brown, subglossy above, matt below; midrib on both surfaces 5-10% covered with fine white-translucent

simple or 3-5-armed stellate hairs, on the upper surface 0.2-0.3 mm diam., on the lower surface

0.1-0.2 mm diam., the leaf-blade otherwise glabrous, apart from sessile red-black glands as the stem,

0.05-0.1 mm diam., 2-6 per mm 2 . Petiole evenly winged along its length, the wings incurved (field

notes); (2.5-)3—4.5 x 0.2-0.4 cm; clasping the stem for l A its circumference, very shortly decurrent by

17

European Journal of Taxonomy 69 : 1-23 ( 2013 )

1-2 mm Lower and intermediate pitchers unknown. Upper pitcher (tendril coiled) 12-15 x 4.5-5 cm;

ovoid-ellipsoid in the lower half, upper half cylindrical, 3-3.5 cm diam., not constricted at any point;

outer surface 10-30% covered in minute red stellate hairs, hairs ca. 0.1 mm diam., both sessile and

shortly stalked, 4-6-armed, arms suberect or patent, density 3-5 per mm 2 , mixed with sessile red-black

glands 0.05 mm diam. as the leaf-blade and stem, hairs denser on lid, and towards the peristome where

they are mixed with sparse erect bushy-bristle hairs 0.2-0.3 mm long; “almost uniformly green with a

few purple spots mainly on the ventricose base” ( Argent et al. 99214); fringed wings are absent, reduced

to inconspicuous ridges; mouth ovate, 4—4.5 x 3-3.5 cm, oblique, slightly concave, “glaucous green

inside with just a few red spots”; peristome (1—)2—3(—5) mm wide, narrowly subcylindrical, rounded

at the front, becoming slightly flattened and widest at the sides, towards the lid, ca. 4 ridges per mm,

ridges 0.075-0.15 mm high, inner edge inconspicuous, holes and teeth not visible (unless dissected: Fig.

4P); outer edge not lobed; column weakly developed, ca. 7 x 3 mm. Ljfl ovate 3.2 x 2.9(-3.2) cm; apex

shallowly retuse, the sinus 3-7 mm wide; base cordate, the sinus 4 mm deep, 8-15 mm wide; green;

margin undulate; lower surface with convex basal appendage, 0.4-0.7 x 1-2 mm, arising from near the

midpoint of the 5-6 x 0.5 mm long basal midline ridge; nectar glands slightly dimorphic, each with a

different distribution: (type 1) moderately dense on the basal ridge and appendage (Fig. 4L) and in a ca.

2 mm band each side (but not extending along midline), glands with raised borders, shortly elliptic, 0.1—

0.2(-0.3) mm long; (type 2) slightly larger, (0. l-)0.2-0.3 mm long, moderately dense, in bands 2-4 mm

wide along the lid margins, 25—40 glands on each side, one sheet (atypical?) with a few additional large

elliptic glands, 0.7 x 0.4 mm, bordered, very sparsely scattered between the margins; sessile red-black

glands, as stem, leaf and outer pitcher surface, 0.005-0.01 mm diam., scattered over surface ca. 3 per

mm 2 ; marginal part of lower surface with a few minute stellate hairs. Spur inserted 2 mm below junction

of lid and pitcher on ridge; simple, stout at base tapering to a long, acute apex; 7-9.5 x 0.5-0.7 mm;

completely covered in long, grey appressed hairs, hairs (0.5—)0.7—1(—1.2) mm long. Upper surface of lid

with two prominent nerves, nerves densely (80-90% cover) white hairy, hairs of two types: (1) basally

1-2-branched hairs 0.3-0.4 mm long, (2) minute 3-5-armed stellate hairs 0.1 mm diam.; remainder of

lid surface with type (2) hairs, but indumentum 30M0% cover, and with sparse perithecoid nectar glands

0.25 mm long. Inflorescence and infructescence unknown.

Distribution & habitat

Philippines, Visayas, Leyte; volcanic geology; “climbing on fallen tree in submontane mossy forest”,

elevation 900 m ( Argent et al. 99214).

Conservation

Nepenthes leyte sp. nov. is known currently from a single individual in an unprotected area, in a country,

including specifically the island of Leyte, where most of the original forest habitat has been cleared

for timber and agricultural land and where forest degradation and clearance are ongoing (Myers et al.

2000; GoogleEarth viewed 2 Oct. 2013). Accordingly, it is here assessed as Critically Endangered under

Criterion D of IUCN (2012). It is to be hoped that further exploration will reveal additional localities for

the species, and that protection can be arranged before it becomes extinct. Currently no protected areas

are believed to be present on Leyte apart from the 2193 ha Lake Danao National or Natural Park which

is about 14 km to the W of the type location. However this seems to be mainly a recreational area, and

within the reserve, illegal settlement, slash and burn agriculture and illegal logging are reported to be

problems ( http://en,wikipedia.org/wiki/Lake_Danao_(Levte)#Threats . viewed 12 Oct. 2013). Viewing

the area immediately around Lake Danao on GoogleEarth (2007 imagery, viewed 12 Oct 2013) confirms

that large areas have been and were in 2007 in the process of being cleared and inhabited, and that

these activities extend towards the E and the only known location for N. leyte sp. nov. Some original

forest still survives along the central high ridge of Leyte, where terrain appears rugged, including the

location indicated as the type location of the species, however the resolution of the imagery here is not

18

CHEEK M. & JEBB M., Nepenthes (Caryophyllales: Nepenthaceae) in the Philippines

Fig. 4. Nepenthes leyte sp. nov. A. Habit, climbing stem with upper pitchers. B. Stem section showing

axillary hair patch and supra-axillary bud. C. Lower surface of leaf-blade, with sessile glands. D. Midrib

of leaf-blade, lower surface, stellate to simple hairs. E. Midrib of leaf-blade, upper surface with stellate

hairs. F. Upper lid of pitcher, upper surface. G. Indumentum of upper surface of lid, over nerve. H. As

G but distant from nerve. I. Spur of upper pitcher and junction of lid with peristome (inverted). J. Lid of

upper pitcher, lower surface. K. Detail of J showing large nectar glands. L. Detail of J showing midline

ridge with appendage and type (1) nectar glands. M. Indumentum of outer pitcher surface. N. Peristome

of upper pitcher viewed from above. O. Peristome viewed from inside pitcher. P. Peristome dissected

to expose the inner edge, with teeth. Q. Peristome, transverse section (outer surface on right). All drawn

from Argent et al. 99214 by Andrew Brown.

19

European Journal of Taxonomy 69 : 1-23 ( 2013 )

sufficiently high to gauge how intact the habitat is. The eastern side of Leyte has higher rainfall and

the forest has extensively been replaced by intensive industrial oil palm plantations which extended in

2003 to within 4 km of the type location (GoogleEarth imagery dating from 2003, viewed 2 Oct. 2013).

Collection of Nepenthes leyte sp. nov. from the wild to supply the horticultural trade is considered a low

risk for this species since its pitchers are not as spectacular or as bizarre as those of other members of

the genus in the Philippines.

Remarks

Argent et al. 99214 , here described as Nepenthes leyte sp. nov., while superficially similar to N.

graciliflora , the only other species of the genus known on Leyte ( Wenzel 680 , GH!; Barbon et al. in PPI

8735 , BRIT!; ibid. 8561 , BRIT!), cannot be confused with it. This is due to the stellate hairs present on

the outer pitcher surface of Nepenthes leyte sp. nov. (versus absent in N. graciliflora :), and the dimorphic

nectar lid glands concentrated around the margin and appendage (not monomorphic, uniformly dense

and distributed). It also has petioles that appear cylindrical since the wings are involute (not with

patent wings). Apart from N. graciliflora itself, only one other member of the N. alata group is, so far,

known from the Visayas: N. negros (Negros and Biliran islands). However, N. negros has densely hairy

stems (versus glabrous), upper pitcher with fringed wings (not absent) and the inner peristome edge

has conspicuous teeth and holes (versus conspicuous teeth absent). Nepenthes leyte sp. nov. can be

distinguished from other species of the Nepenthes alata group using the key above.

Discussion

In an earlier paper, the Nepenthes alata group of species, confined to the Philippines, was characterised

and a key was provided to the then recognised species (Cheek & Jebb 2013d). In that paper the species

of the N. alata group were characterised as possessing: (1) a basal appendage on the lower surface of the

pitcher lid, (2) a terete stem, (3) a distinct, but winged petiole, the petiole wings wide, decurrent from

the blade, (4) the peristome finely ridged, the outer edge not or only slightly lobed, (5) the inner surface

lacking conspicuous teeth, (6) the mouth ovate, oblique, without a well-developed column (Cheek &

Jebb 2013d).

The four new species described here have characters that necessitate modification of the Nepenthes

alata group as circumscribed in Cheek & Jebb (2013d). All but N. leyte sp. nov. of the new species

have stems that can be 4-angled in section rather than terete. In N. extincta sp. nov. and in N. leyte

sp. nov., the petiole wings are incurved, the two edges overlapping, giving the petiole a cylindrical

appearance. Both N. extincta sp. nov. and N. kitanglad sp. nov. have broad, partly flattened peristomes

(rather than narrowly cylindric) and pitcher mouths which are concave (rather than straight). In the case

of N. kitanglad sp. nov., a moderately well-developed column is formed (rather than none, or a weakly

developed column). Several of these characters are more usual in the Regiae group of species of Danser

(1928), which are considered by us to be closely related to the N. alata group. The two species groups

share (1) petiolate leaves and, (2) on the basal part of the lower surface of the lid, appendages emerging

from a keel-lilce ridge or crest, and (3) inflorescences with 2-flowered partial-peduncles. The two species

groups can be both defined and separated from each other using the key below. It is possible that future

discoveries will provide evidence that the two groups are a continuum, and should be united.

- Upper pitchers infundibuliform or infundibuliform-cylindrical, always widest at the peristome;

petioles not or weakly winged, U-shaped in section, the wing width if present, shorter than the width

of the central, non-winged part of the petiole; stem and leaf indumentum of persistent, red-brown

bristle-like hairs. BORNEO, SULAWESI & NEW GUINEA.

. Regiae species group (Danser 1928)

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CHEEK M. & JEBB M., Nepenthes (Caryophyllales: Nepenthaceae) in the Philippines

- Upper pitchers usually ovoid-cylindrical, rarely subcylindrical, always widest in the basal half

(except in N. ceciliae and N. copelandii ); petiole wing width wider than the width of the central,

non-winged part of the petiole, wings either held flat or inrolled and then the petiole

appearing cylindrical. PHILIPPINES EXCLUDING PALAWAN.

. N. alata species group (Cheek & Jebb 2013d)

Acknowledgements

Janis Shillito typed the manuscript. Barbara Kennedy at BISH and Tiana Rehman at BRIT kindly

provided the loan of the material cited. Lee Davies and Yvette Harvey digitised the images.

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Manuscript received: 23 September 2013

Manuscript accepted: 19 November 2013

Published on: 12 December 2013

Topic editor: Thomas Janssen

Desk editor: Natacha Beau

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; National Botanic Garden

of Belgium, Meise, Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History

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Natural history Museum of Denmark, Copenhagen, De nm ark

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