''Nepenthes macrophylla'' , the large-leaved pitcher-plant,Phillipps, A. & A. Lamb 1996. ''

Pitcher-Plants of Borneo ''Pitcher-Plants of Borneo'' is a monograph by Anthea Phillipps and Anthony Lamb on the tropical pitcher plants of Borneo. It was first published in 1996 by Natural History Publications (Borneo), in association with the Royal Botanic Gardens, ...

''. Natural History Publications (Borneo), Kota Kinabalu. is a tropical

pitcher plant Pitcher plants are several different carnivorous plants which have modified leaves known as pitfall traps—a prey-trapping mechanism featuring a deep cavity filled with digestive liquid. The traps of what are considered to be "true" pitcher p ...

known only from a very restrictive elevation on

Mount Trusmadi Mount Trusmadi or Trus Madi ( ms, Gunung Trusmadi) is a mountain located at the Interior Division of Sabah, Malaysia. It is considered as the second highest mountain in both Sabah and Malaysia at , after Mount Kinabalu with Trusmadi offering a ...

Sabah Sabah () is a state of Malaysia located in northern Borneo, in the region of East Malaysia. Sabah borders the Malaysian state of Sarawak to the southwest and the North Kalimantan province of Indonesia to the south. The Federal Territory o ...

Malaysian Borneo East Malaysia (), or the Borneo States, also known as Malaysian Borneo, is the part of Malaysia on and near the island of Borneo, the world's third-largest island. Near the coast of Sabah is a small archipelago called Labuan. East Malaysia li ...

. ''Nepenthes macrophylla'' was once thought to be a subspecies of ', but differs from that species in several aspects of pitcher and leaf

morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...

.Clarke, C.M. 1997. ''

Nepenthes of Borneo ''Nepenthes of Borneo'' is a monograph by Charles Clarke on the tropical pitcher plants of Borneo.Clarke, C.M. 1997. ''Nepenthes of Borneo''. Natural History Publications (Borneo), Kota Kinabalu. It was first published in 1997 by Natural History ...

''. Natural History Publications (Borneo), Kota Kinabalu.

Botanical history

''Nepenthes macrophylla'' was known to grow on

for a long time prior to its description, although it was not initially considered a distinct species. Shortly after its discovery, it was lumped with ', a similar species from

Kinabalu National Park Kinabalu Park ( ms, Taman Kinabalu), established as one of the first national parks of Malaysia in 1964, is Malaysia's first World Heritage Site designated by UNESCO in December 2000 for its "outstanding universal values" and the role as one of ...

. In his 1976 guide, ''

Nepenthes of Mount Kinabalu ''Nepenthes of Mount Kinabalu'' is a monograph by Shigeo Kurata on the tropical pitcher plants of Mount Kinabalu and the surrounding area of Kinabalu National Park in Sabah, Borneo. It was published in 1976 by Sabah National Parks Trustees as th ...

'', botanist

Shigeo Kurata is a Japanese botanist and '' Nepenthes'' taxonomist whose work in the 1960s and 1970s contributed much to the current popularity of these plants.Clarke, C.M. & C.C. Lee 2004. ''Pitcher Plants of Sarawak''. Natural History Publications (Borneo), ...

wrote:Kurata, S. 1976. ''

''. Sabah National Parks Publications No. 2, Sabah National Parks Trustees, Kota Kinabalu.

Mt. Trus Madi, located in the south of Mt. Kinabalu, become known as a locality for this species Though this information is not confirmed yet, its existence is expected because this mountain falls within the vertical distribution of this species.

Later, ''N. macrophylla'' was treated as a form of ', another closely related species.Marabini, J. 1984. ''

Carnivorous Plant Newsletter The ''Carnivorous Plant Newsletter'' is the official publication of the International Carnivorous Plant Society (ICPS), the largest such organization in the world. It is headquartered in Walnut Creek, California. History and editorship The newsle ...

'' 13(2): 38–40. Phillipps, A., A. Lamb & C.C. Lee 2008. ''

Pitcher Plants of Borneo ''Pitcher-Plants of Borneo'' is a monograph by Anthea Phillipps and Anthony Lamb on the tropical pitcher plants of Borneo. It was first published in 1996 by Natural History Publications (Borneo), in association with the Royal Botanic Gardens, Kew ...

''. Second Edition. Natural History Publications (Borneo), Kota Kinabalu. An early collection of ''N. macrophylla'' was made by Johannes Marabini in March 1983 and the herbarium material, designated as ''Marabini 2167'',Schlauer, J. N.d
''Nepenthes macrophylla''
Carnivorous Plant Database. deposited at the

University of Erlangen-Nuremberg A university () is an institution of higher (or tertiary) education and research which awards academic degrees in several academic disciplines. Universities typically offer both undergraduate and postgraduate programs. In the United States, ...

Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...

. Four years later, Marabini formally described the

taxon In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...

as a subspecies of ''N. edwardsiana''. In their 1996 monograph, ''

'', Anthea Phillipps and Anthony Lamb treated ''N. macrophylla'' as a species in the process of being described, referring to it as "''Nepenthes macrophylla'' (Marabini) Jebb & Cheek ined.". Botanists

Matthew Jebb Matthew Hilary Peter Jebb (born 1958) is an Irish botanist and taxonomist specialising in the ant plant genera ''Squamellaria'', '' Myrmecodia'', '' Hydnophytum'', ''Myrmephytum'' and '' Anthorrhiza'', as well as the carnivorous plant genus '' ...

and

Martin Cheek Martin Roy Cheek (born 1960) is a botanist and taxonomist at the Royal Botanic Gardens, Kew.A skeletal revision of ''Nepenthes'' (Nepenthaceae)" in the botanical journal ''

Blumea ''Blumea'' is a genus of flowering plants of the family Asteraceae. Characteristics Genus ''Blumea'' is found in the tropical and sub-tropical zones of Asia, especially the Indian Subcontinent and Southeast Asia. A few species are found in Aust ...

''. This interpretation was supported by

Charles Clarke Charles Rodway Clarke (born 21 September 1950) is a British Labour Party politician, who was the Member of Parliament (MP) for Norwich South from 1997 until 2010, and served as Home Secretary from December 2004 until May 2006. Early life T ...

, who noted that ''N. edwardsiana'' and ''N. villosa'' "have more in common with each other than do ''N. edwardsiana'' and ''N. macrophylla''".

Description

''Nepenthes macrophylla'' is a climbing plant. The stem reaches a length of more than 10 m and is up to 10 mm in diameter. Internodes are up to 35 cm long and circular in cross section. Leaves are

coriaceous This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...

and petiolate. The

lamina Lamina may refer to: Science and technology * Planar lamina, a two-dimensional planar closed surface with mass and density, in mathematics * Laminar flow, (or streamline flow) occurs when a fluid flows in parallel layers, with no disruption betwee ...

or leaf blade is oblong in shape and reaches exceptionally large dimensions of up to 60 cm by 20 cm. The apex of the lamina is

acuminate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular ...

-obtuse, while the base is abruptly contracted where the petiole begins. The petiole itself is up to 15 cm long. It is canaliculate (grooved lengthwise) and typically bears wings that form a partially amplexicaul sheath around the stem. One to three longitudinal veins are present on either side of the

midrib This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...

, but

pinnate Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and in ...

veins are indistinct.

Tendril In botany, a tendril is a specialized stem, leaf or petiole with a threadlike shape used by climbing plants for support and attachment, as well as cellular invasion by parasitic plants such as '' Cuscuta''. There are many plants that have tend ...

s measure up to 35 cm in length. The pitchers of ''N. macrophylla'' are robust and almost woody in texture. Rosette and lower pitchers are shortly cylindrical in shape, becoming narrower around the midpoint. They can be very large: up to 35 cm high by 15 cm wide. A pair of fringed wings (≤10 mm wide) runs down the ventral surface of the pitcher. Terrestrial pitchers usually have a hip in their lower half. The pitcher mouth is

ovate Ovate may refer to: * Ovate (egg-shaped) leaves, tepals, or other botanical parts *Ovate, a type of prehistoric stone hand axe *Ovates, one of three ranks of membership in the Welsh Gorsedd *Vates In modern English, the nouns vates () and ova ...

and has an oblique insertion. The

peristome Peristome (from the Greek ''peri'', meaning 'around' or 'about', and ''stoma'', 'mouth') is an anatomical feature that surrounds an opening to an organ or structure. Some plants, fungi, and shelled gastropods have peristomes. In mosses In mosse ...

is cylindrical and expanded, measuring up to 15 mm wide. It bears a series of highly developed ribs, which terminate in prominent downward-pointing teeth. The pitcher lid or operculum is ovate, very broad, and has a

cordate Cordate is an adjective meaning ' heart-shaped' and is most typically used for: * Cordate (leaf shape) The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a ...

base. An unbranched

spur A spur is a metal tool designed to be worn in pairs on the heels of riding boots for the purpose of directing a horse or other animal to move forward or laterally while riding. It is usually used to refine the riding aids (commands) and to ba ...

(≤9 mm long) is inserted near the base of the lid. Upper pitchers are similar to their lower counterparts in most respects. They differ in having a more cylindrical shape, reduced wings, and the hip always being located in the upper half. Traps may have a range of colours, although the yellow-pitchered form is more common than the red-pitchered form. ''Nepenthes macrophylla'' has a

racemose A raceme ( or ) or racemoid is an unbranched, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are produced as the s ...

inflorescence An inflorescence is a group or cluster of flowers arranged on a Plant stem, stem that is composed of a main branch or a complicated arrangement of branches. Morphology (biology), Morphologically, it is the modified part of the shoot of sperma ...

, with both male and female ones having a similar structure. The peduncle is up to 23 cm long and 5 mm wide, while the

rachis In biology, a rachis (from the grc, ῥάχις [], "backbone, spine") is a main axis or "shaft". In zoology and microbiology In vertebrates, ''rachis'' can refer to the series of articulated vertebrae, which encase the spinal cord. In this c ...

is up to 55 cm long.

Pedicels In botany, a pedicel is a stem that attaches a single flower to the inflorescence. Such inflorescences are described as ''pedicellate''. Description Pedicel refers to a structure connecting a single flower to its inflorescence. In the absenc ...

are one-flowered, bracteate, and measure up to 16 mm in length.

Sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coine ...

s are elliptic in shape and up to 6 mm long. Most parts of the plants bear an

indumentum In biology, an indumentum (Latin, literally: "garment") is a covering of trichomes (fine "hairs") on a plant Davis, Peter Hadland and Heywood, Vernon Hilton (1963) ''Principles of angiosperm taxonomy'' Van Nostrandpage, Princeton, New Jersey, pa ...

of very short hairs, although much of this covering is

caducous Dehiscence is the splitting of a mature plant structure along a built-in line of weakness to release its contents. This is common among fruits, anthers and sporangia. Sometimes this involves the complete detachment of a part; structures that op ...

Ecology

''Nepenthes macrophylla'' is known with certainty only from the summit area of

Borneo Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java, west of Sulawesi, and ea ...

, despite the presence of a number of nearby peaks (such as Mount Pinesowitan) that fall within its altitudinal range. However, there are unconfirmed reports that it also grows "on the summit of a neighbouring peak". Older sources list a number of different values for the altitudinal distribution of ''N. macrophylla'', with some giving a narrow range of 2200–2400 m and others a much wider span of 2000–2600 m. It is now known that this species occurs between 2200 m and the summit at 2642 m.McPherson, S.R. 2009. ''

Pitcher Plants of the Old World ''Pitcher Plants of the Old World'' is a two-volume monograph by Stewart McPherson on the pitcher plants of the genera '' Nepenthes'' and ''Cephalotus''. It was published in May 2009 by Redfern Natural History Productions and covers all species ...

''. 2 volumes. Redfern Natural History Productions, Poole. The typical habitat of ''N. macrophylla'' is mossy

montane forest Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures fall as elevation increases, causing the ecosystem to stratify. This stratification is a crucial ...

along ridge tops, where the vegetation is dominated by '' Leptospermum'' and '' Rhododendron''.Steiner, H. 2002. '' Borneo: Its Mountains and Lowlands with their Pitcher Plants''. Toihaan Publishing Company, Kota Kinabalu. Like most highland ''Nepenthes'' species, ''N. macrophylla'' experiences marked temperature fluctuations between day and night, from a noon maximum of around 30 °C to a nighttime minimum of 5 °C.

Relative humidity Humidity is the concentration of water vapor present in the air. Water vapor, the gaseous state of water, is generally invisible to the human eye. Humidity indicates the likelihood for precipitation, dew, or fog to be present. Humidity dep ...

is always high and in the late evening the summit of Mount Trusmadi is often shrouded in mist and clouds. ''Nepenthes macrophylla'' occurs both terrestrially and as an epiphyte. The species is sympatric with ' and '. A

natural hybrid In biology, a hybrid is the offspring resulting from combining the qualities of two organisms of different breeds, varieties, species or genera through sexual reproduction. Hybrids are not always intermediates between their parents (such as in ...

between ''N. lowii'' and ''N. macrophylla'', described as ', has been recorded. Due to its restricted natural range and the fact that Mount Trusmadi does not receive the same high level of protection as

Mount Kinabalu Mount Kinabalu ( ms, Gunung Kinabalu, Dusun: ''Gayo Ngaran or Nulu Nabalu'') is the highest mountain in Borneo and Malaysia. With an elevation of , it is third-highest peak of an island on Earth, and 20th most prominent mountain in the worl ...

, ''N. macrophylla'' is classified as Critically Endangered on the 2006 IUCN Red List of Threatened Species. This differs slightly from an informal assessment made by botanist

in 1997, who classified the species as

Endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and in ...

based on the IUCN criteria. Clarke wrote that the species "requires the highest level of protection if it is to persist in the wild into the next century". ''N. macrophylla'' is threatened by over-collection and damage caused to its habitat by visitors. These activities have already depleted populations of this species on the mountain and it is thought to be the most endangered highland ''Nepenthes'' of Borneo.

Carnivory

''Nepenthes macrophylla'' has formed a mutualistic relationship with the mountain treeshrew (''Tupaia montana''). The pitchers of ''N. macrophylla'' exude a sugary reward on the reflexed pitcher lid and provide a perch for the visitor. The treeshrews eat this exudate and defecate into the pitchers, supplying the plant with valuable

nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...

. A 2010 study showed that the shape and size of the pitcher orifice of ''N. macrophylla'' exactly match the dimensions of a typical ''Tupaia montana''.Chin, L., J.A. Moran & C. Clarke 2010. Trap geometry in three giant montane pitcher plant species from Borneo is a function of tree shrew body size. ''New Phytologist'' 186 (2): 461–470. Walker, M. 2010
Giant meat-eating plants prefer to eat tree shrew poo
''BBC Earth News'', March 10, 2010. A similar adaptation was found in '' N. lowii'' and '' N. rajah'', and is also likely to be present in '' N. ephippiata''.Clarke, C., J.A. Moran & L. Chin 2010. Mutualism between tree shrews and pitcher plants: perspectives and avenues for future research. ''Plant Signaling & Behavior'' 5(10): 1187–1189. Greenwood, M., C. Clarke, C.C. Lee, A. Gunsalam & R.H. Clarke 2011. A unique resource mutualism between the giant Bornean pitcher plant, ''Nepenthes rajah'', and members of a small mammal community. ''PLoS ONE'' 6(6): e21114. In all three of these species (''N. ephippiata'' has not been investigated), the colour of the lower lid surface corresponds to ''T. montana'' visual sensitivity maxima in the green and blue wavebands, making it stand out against adjacent parts of the pitcher.

Related species

''Nepenthes macrophylla'' is most closely related to ' and '. In his Carnivorous Plant Database, taxonomist

Jan Schlauer Jan, JaN or JAN may refer to: Acronyms * Jackson, Mississippi (Amtrak station), US, Amtrak station code JAN * Jackson-Evers International Airport, Mississippi, US, IATA code * Jabhat al-Nusra (JaN), a Syrian militant group * Japanese Article Numb ...

treats ''N. macrophylla'' as a possible synonym of ''N. villosa''. ''Nepenthes macrophylla'' differs from both of these species in the structure of its peristome. Although highly developed, the peristome ribs and teeth of ''N. macrophylla'' are considerably shorter and more widely spaced than those of either ''N. edwardsiana'' or ''N. villosa''. The pitcher mouth of ''N. macrophylla'' is distinctive in that it rises gradually towards the lid, while at the same time not forming a pronounced neck. In addition, the mouth of this species has a much more oblique insertion than its relatives. ''Nepenthes macrophylla'' is also distinguished by its broad,

lid. The lower pitchers of ''N. edwardsiana'' and ''N. macrophylla'' are quite similar in shape, although in the latter species the hip is always positioned in the upper portion of the pitcher cup. The upper pitchers of these species are more distinct, with those of ''N. macrophylla'' being more ovoid and less elongated. As its name suggests, ''N. macrophylla'' produces very large leaves and these may be twice as long as those of either ''N. edwardsiana'' or ''N. villosa''. Whereas ''N. edwardsiana'' and ''N. villosa'' are restricted to generally

ultramafic Ultramafic rocks (also referred to as ultrabasic rocks, although the terms are not wholly equivalent) are igneous and meta-igneous rocks with a very low silica content (less than 45%), generally >18% MgO, high FeO, low potassium, and are composed ...

soils in the Kinabalu area, ''N. macrophylla'' is only found near the summit of

, which is mostly composed of

sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates ...

. Botanists

and

Martin Cheek Martin Roy Cheek (born 1960) is a botanist and taxonomist at the Royal Botanic Gardens, Kew.endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...

Palawan Palawan (), officially the Province of Palawan ( cyo, Probinsya i'ang Palawan; tl, Lalawigan ng Palawan), is an archipelagic province of the Philippines that is located in the region of Mimaropa. It is the largest province in the country in t ...

in the

Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no), * bik, Republika kan Filipinas * ceb, Republika sa Pilipinas * cbk, República de Filipinas * hil, Republ ...

.Schlauer, J. 2000. ''

'' 29(2): 53.

Natural hybrids

In the wild, ''N. macrophylla'' is only known to hybridise with '. ''N. lowii'' × ''N. macrophylla'' was discovered on

by Johannes Marabini and John Briggs in 1983. Later that year, it was described as by Marabini. Briggs returned to Mount Trusmadi in 1984, but found only one small group of plants. has petiolate leaves measuring up to 50 cm in length. The pitchers of this hybrid are some of the largest of any Bornean ''Nepenthes'' species,Briggs, J.G.R. 1988. ''Mountains of Malaysia: A Practical Guide and Manual''. Longman Malaysia, Kuala Lumpur. reaching 35 cm in height. They are roughly intermediate in form between those of its parent species. The lid is held away from the mouth as in ''N. lowii'' and bears short bristles on its lower surface. The peristome has prominent ribs and teeth, but is not as developed as that of ''N. macrophylla''.Clarke, C.M. 2001. ''

A Guide to the Pitcher Plants of Sabah This list of ''Nepenthes'' literature is a listing of major published works dealing with the tropical pitcher plants of the genus ''Nepenthes''. It includes specialised standalone publications and Biological classification, taxonomic monographs ...

''. Natural History Publications (Borneo), Kota Kinabalu. The

of may be up to 50 cm long and has two-flowered

pedicels In botany, a pedicel is a stem that attaches a single flower to the inflorescence. Such inflorescences are described as ''pedicellate''. Description Pedicel refers to a structure connecting a single flower to its inflorescence. In the absenc ...

. Despite the size of the pitchers, this hybrid is not large in stature. is restricted to the summit ridge of Mount Trusmadi and has been recorded from elevations of 2500 to 2600 m above sea level.

Notes

:a.The pitchers of ''N. macrophylla'' have been described as having "the consistency of cardboard" and because of this they find use as bottles among farmers from

Tambunan Tambunan ( ms, Pekan Tambunan) is the capital of the Tambunan District in the Interior Division of Sabah, Malaysia. Its population was estimated to be around 35,667 in 2010. It is located 80 kilometres east of the state capital, Kota Kinabalu ...

References

External links

*
Nepenthes macrophylla
' at

ARKive ARKive was a global initiative with the mission of "promoting the conservation of the world's threatened species, through the power of wildlife imagery", which it did by locating and gathering films, photographs and audio recordings of the worl ...

Video of tree shrew visiting pitchers of ''N. macrophylla''
{{Taxonbar, from=Q146125 Carnivorous plants of Asia Coprophagous plants macrophylla Endemic flora of Borneo Flora of Sabah Critically endangered plants Plants described in 1997