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''Nepenthes aristolochioides'' 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 ...
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 elsew ...
to
Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
, where it grows at elevations of 1800–2500 m above sea level. It has an extremely unusual pitcher morphology, having an almost vertical opening to its traps.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 kno ...
''. 2 volumes. Redfern Natural History Productions, Poole.
It is critically endangered by overcollection. The
specific epithet In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
''aristolochioides'' is formed from the genus name ''
Aristolochia ''Aristolochia'' () is a large plant genus with over 500 species that is the type genus of the family Aristolochiaceae. Its members are commonly known as birthwort, pipevine or Dutchman's pipe and are widespread and occur in the most diverse clima ...
'' and the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
ending ''-oides'', meaning "resembling". It refers to the similarity that the pitchers of this species bear, in both shape and pigmentation, to the specialised flowers of ''Aristolochia''.


Botanical history

''Nepenthes aristolochioides'' was first collected by
Willem Meijer Willem Meijer (1923 – 22 October 2003) was a Dutch botanist and plant collector. Background and education Meijer was born in 1923 in The Hague, Netherlands. He received his Ph.D. from the University of Amsterdam in 1951. Meijer travelled to J ...
on August 5, 1956. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
, ''Meijer 6542'', was collected on that date from
Mount Tujuh Mount Tujuh ( id, Gunung Tujuh) is a caldera volcano in the Barisan Mountains of Sumatra. It has seven (tujuh) peaks, of which only three have been climbed.Clarke, C.M. 2001. ''Nepenthes of Sumatra and Peninsular Malaysia''. Natural History Public ...
(Tudjuh) in
Jambi Jambi is a province of Indonesia. It is located on the east coast of central Sumatra and spans to the Barisan Mountains in the west. Its capital and largest city is Jambi. The province has a land area of 50,160.05 km2, and a sea area of 3, ...
at an elevation of 2000 m. It is deposited at the
National Herbarium of the Netherlands National may refer to: Common uses * Nation or country ** Nationality – a ''national'' is a person who is subject to a nation, regardless of whether the person has full rights as a citizen Places in the United States * National, Maryland, ce ...
(L) in
Leiden Leiden (; in English and archaic Dutch also Leyden) is a city and municipality in the province of South Holland, Netherlands. The municipality of Leiden has a population of 119,713, but the city forms one densely connected agglomeration wit ...
, but is in relatively poor condition.Nerz, J. 1998
Rediscovery of an outstanding ''Nepenthes'': ''N. aristolochioides'' (Nepenthaceae)
''
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 ...
'' 27(3): 101–114.
An isotype is held at Herbarium Bogoriense (BO), the herbarium of the
Bogor Botanical Gardens The Bogor Botanical Gardens ( id, Kebun Raya Bogor) is a botanical garden located in Bogor, Indonesia, 60 km south of central Jakarta. It is currently operated by Indonesian Institute of Sciences (Indonesian: ''Lembaga Ilmu Pengetahuan Indo ...
(formerly the Herbarium of the Buitenzorg Botanic Gardens) in
Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's List ...
.Clarke, C.M. 2001. ''
Nepenthes of Sumatra and Peninsular Malaysia ''Nepenthes of Sumatra and Peninsular Malaysia'' is a monograph by Charles Clarke on the tropical pitcher plants of Sumatra, Peninsular Malaysia, and their minor surrounding islands. It was published in 2001 by Natural History Publications (Borneo ...
''. Natural History Publications (Borneo), Kota Kinabalu.
Although labelled as "new species?", the specimen was largely overlooked for over 30 years. In 1988, botanist
Joachim Nerz Dr. Joachim Nerz (born 1964) is a German taxonomist and botanist specialising in the carnivorous plant genera ''Heliamphora'' and '' Nepenthes''. Nerz has described several new species, mostly with Andreas Wistuba. Publications * Schlauer, J. ...
became aware of it upon visiting the herbarium of
Leiden University Leiden University (abbreviated as ''LEI''; nl, Universiteit Leiden) is a Public university, public research university in Leiden, Netherlands. The university was founded as a Protestant university in 1575 by William the Silent, William, Prince o ...
. The name ''N. aristolochioides'' was coined quite early on; it was already being used in 1994 to informally refer to this (at the time undescribed)
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 nam ...
. That same year, 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 ...
questioned the supposed lateral pitcher mouth of ''N. aristolochioides'' in email correspondence with botanist
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 ''Nepe ...
, who was preparing a revision of the genus at the time.Schlauer, J. 1994
NEPENTHES comments for Flora Malesiana
Carnivorous Plant Mailing List, September 9, 1994.
Schlauer wrote that he had examined a specimen of this species (''Meijer 7426'') and that the seemingly vertical insertion of the pitcher mouth might be a result of the preservation process, whereby the traps had become "compressed along their longitudinal axes". In the summer of 1996, Nerz met with Schlauer and Meijer in the
Frankfurt Frankfurt, officially Frankfurt am Main (; Hessian: , "Frank ford on the Main"), is the most populous city in the German state of Hesse. Its 791,000 inhabitants as of 2022 make it the fifth-most populous city in Germany. Located on its na ...
Palmengarten The Palmengarten is one of three botanical gardens in Frankfurt am Main, Germany. It is located in the Westend (Frankfurt am Main), Westend-Süd district. It covers a surface of 22 hectares. It is a major tourist attraction. History Like ma ...
, where Meijer showed him a photograph of the mysterious species. Together with Katrin Hinderhofer, Nerz organised a field trip to Sumatra in June 1996 and was successful in rediscovering ''N. aristolochioides'' in the wild. ''Nepenthes aristolochioides'' was finally described by
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 ''Nepe ...
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)", published in the May 1997 issue of 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 ...
''.''Nepenthes aristolochioides'' M.Jebb & Cheek
International Plant Names Index The International Plant Names Index (IPNI) describes itself as "a database of the names and associated basic bibliographical details of seed plants, ferns and lycophytes." Coverage of plant names is best at the rank of species and genus. It inclu ...
(IPNI).
Joachim Nerz Dr. Joachim Nerz (born 1964) is a German taxonomist and botanist specialising in the carnivorous plant genera ''Heliamphora'' and '' Nepenthes''. Nerz has described several new species, mostly with Andreas Wistuba. Publications * Schlauer, J. ...
wrote a detailed description of the species for an issue of the ''
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 ...
'' published the following year. The next major treatments of the species appeared in Cheek and Jebb's updated 2001 work, "
Nepenthaceae ''Nepenthes'' () is a genus of carnivorous plants, also known as tropical pitcher plants, or monkey cups, in the monotypic family Nepenthaceae. The genus includes about 170 species, and numerous natural and many cultivated hybrids. They are mos ...
";Cheek, M.R. & M.H.P. Jebb 2001.
Nepenthaceae ''Nepenthes'' () is a genus of carnivorous plants, also known as tropical pitcher plants, or monkey cups, in the monotypic family Nepenthaceae. The genus includes about 170 species, and numerous natural and many cultivated hybrids. They are mos ...
. ''Flora Malesiana'' 15: 1–157.
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 Th ...
's ''
Nepenthes of Sumatra and Peninsular Malaysia ''Nepenthes of Sumatra and Peninsular Malaysia'' is a monograph by Charles Clarke on the tropical pitcher plants of Sumatra, Peninsular Malaysia, and their minor surrounding islands. It was published in 2001 by Natural History Publications (Borneo ...
'', also published in 2001; and Stewart McPherson's two-volume ''
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 kno ...
'', released in 2009, which included colour photographs of specimens from a newly discovered locality.


Description

''Nepenthes aristolochioides'' is a climbing plant. The stem, which may be branched, is
terete Terete is a term in botany used to describe a cross section that is circular, or like a distorted circle, with a single surface wrapping around it.Lichen Vocabulary, Lichens of North America Information, Sylvia and Stephen Sharnoff/ref> This is us ...
and grows to 8 m in length and 5 mm in diameter. Internodes are cylindrical to obtusely angular in cross section and up to 15 cm long.
Axillary bud The axillary bud (or lateral bud) is an embryonic or organogenic shoot located in the axil of a leaf. Each bud has the potential to form shoots, and may be specialized in producing either vegetative shoots (stems and branches) or reproductive sh ...
s are notably conspicuous in this species and are located 1.5–7 mm above the
leaf axil A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
s.


Leaves

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
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
. 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 ...
is linear,
lanceolate 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 o ...
or
spathulate 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 ...
-lanceolate in form and up to 20 cm long by 5 cm wide. It has an acute or obtuse apex that may rarely be sub-
peltate 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 o ...
. It is gradually
attenuate In physics, attenuation (in some contexts, extinction) is the gradual loss of flux intensity through a medium. For instance, dark glasses attenuate sunlight, lead attenuates X-rays, and water and air attenuate both light and sound at variable a ...
towards the base, becoming partly amplexicaul (clasping the stem for one-third to half of its circumference) and, rarely, slightly
decurrent ''Decurrent'' (sometimes decurring) is a term used in botany and mycology to describe plant or fungal parts that extend downward. In botany, the term is most often applied to leaf blades that partly wrap or have wings around the stem or petio ...
. Leaves of short stems have rounded auricles, whereas those of climbing stems lack auricles. Two to five 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 ...
. They arise from the leaf base and occasionally along the length of the midrib, and are restricted to the distal third to two-thirds of the lamina, where they run parallel to the laminar margin. These longitudinal veins are indistinct in dried specimens.
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 irregularly
reticulate Reticulation is a net-like pattern, arrangement, or structure. Reticulation or Reticulated may refer to: * Reticulation (single-access key), a structure of an identification tree, where there are several possible routes to a correct identificatio ...
and indistinct. They are few in number and arise obliquely to eventually curve towards the laminar apex. The lower laminar surface bears sessile glands.
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 tendr ...
s reach 15 cm in length. Those bearing rosette pitchers are typically around twice as long as the laminae and do not have a curl.


Pitchers

''Nepenthes aristolochioides'' is noted for exhibiting relatively little dimorphism between its lower and upper pitchers. Rosette and lower pitchers are only briefly produced on small rosettes before the plant begins to climb, or on offshoots from the climbing stem. They arise from the ends of the tendrils, forming a 3–5 mm wide curve. They are broadly
infundibular An infundibulum (Latin for ''funnel''; plural, ''infundibula'') is a funnel-shaped cavity or organ. Anatomy * Brain: the pituitary stalk, also known as the ''infundibulum'' and ''infundibular stalk'', is the connection between the hypothalamus and ...
in the lower two-thirds and
globose A sphere () is a geometrical object that is a three-dimensional analogue to a two-dimensional circle. A sphere is the set of points that are all at the same distance from a given point in three-dimensional space.. That given point is the ce ...
above, forming a dome above the pitcher opening. They reach 7 cm in height and 3 cm in width. A pair of wings (≤9 mm wide) runs down the front of the pitcher, either covering the length of the trap's ventral surface or being restricted to the upper part only. These wings bear fringe elements up to 10 mm long. The pitcher mouth is
orbicular Orbicular is an adjective meaning "circular" Orbicular may also refer to: * Orbicular leaf, a plant leaf shape * Orbicularis oculi muscle, a muscle around the eye * Orbicularis oris muscle, a muscle around the mouth * Orbicular batfish, a specie ...
to ovate and up to 1.5 cm in diameter. It has a horizontal, oblique or almost vertical insertion. The
gland In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream (endocrine gland) or into cavities inside the body or its outer surface (exocrine gland). Structure De ...
ular region covers almost the entire inner surface of the pitcher, but is often missing from the uppermost parts of the pitcher dome. The waxy zone typical of many ''Nepenthes'' species is absent. Digestive glands are overarched; the lower ones measure 0.2–0.3 mm in diameter and are present at a density of around 200/cm2, whereas the upper ones are smaller and present at a density of around 500/cm2. The flattened
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 mosses, ...
is broad, greatly incurved, and up to 20 mm wide. Its ribs are spaced up to 0.5 mm apart. Its inner margin is lined with small teeth that are curled at their apices and are 2–3 times as long as they are broad. The inner portion of the peristome accounts for around 82% of its total cross-sectional surface length.Bauer, U., C.J. Clemente, T. Renner & W. Federle 2012. Form follows function: morphological diversification and alternative trapping strategies in carnivorous ''Nepenthes'' pitcher plants. ''Journal of Evolutionary Biology'' 25(1): 90–102. The pitcher lid or operculum is
orbicular Orbicular is an adjective meaning "circular" Orbicular may also refer to: * Orbicular leaf, a plant leaf shape * Orbicularis oculi muscle, a muscle around the eye * Orbicularis oris muscle, a muscle around the mouth * Orbicular batfish, a specie ...
-
cordate Cordate is an adjective meaning 'heart-shaped' and is most typically used for: * Cordate (leaf shape), in plants * Cordate axe, a prehistoric stone tool See also * Chordate A chordate () is an animal of the phylum Chordata (). All chordat ...
or ovate, up to 1.5 cm wide, and bears no appendages. Large nectar glands are present on the lid's entire lower surface, particularly around the midline. Three prominent veins are usually present on either side of the lid's midline. A broad and flattened
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 back ...
(≤7 mm long) is inserted at the base of the lid. It has been variously described as either branched or unbranched (simple). Upper pitchers gradually arise from the ends of the tendrils, forming a 10 mm wide curve. They are narrowly infundibular in the lower half and utriculate above, with a pronounced dome above the pitcher orifice. The
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Th ...
face of the trap is often noticeably flattened and is around 30% thicker than the wall of the translucent dome. Aerial traps are larger than their terrestrial counterparts, reaching 15 cm in height and 8 cm in width. They typically bear ribs in place of wings, although these may not be apparent at all. The pitcher mouth is subapical, ovate to circular, and measures up to 4 cm in diameter. It is angled even more steeply than in lower pitchers and may be positioned almost vertically. The glandular region usually covers the basal two-thirds of the pitcher cup's inner surface, or the pitcher may be wholly glandular. Digestive glands are slightly overarched; those near the bottom are 0.3–0.4 mm in diameter and occur at a density of around 200/cm2, whereas the upper ones are 0.2–0.3 mm in diameter and occur at a density of around 250/cm2. The peristome, which is up to 20 mm wide, is expanded, incurved, and internally flattened, forming an "entrance corridor" similar to a
lobster pot A lobster trap or lobster pot is a portable trap that traps lobsters or crayfish and is used in lobster fishing. In Scotland (chiefly in the north), the word creel is used to refer to a device used to catch lobsters and other crustaceans. A lo ...
.Rice, B. 2007. Carnivorous plants with hybrid trapping strategies. ''
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 ...
'' 36(1): 23–27.
It extends for up to 2 mm on its outer margin, which is rounded. There is often a gap of up to 5 mm between the two lobes of the peristome near the base of the lid. The peristome bears ribs up to 0.8 mm high and spaced up to 0.8 mm apart, which terminate in indistinct teeth that are 2–3 times as long as they are broad. Large nectaries are located between the ribs. The pitcher lid is
orbicular Orbicular is an adjective meaning "circular" Orbicular may also refer to: * Orbicular leaf, a plant leaf shape * Orbicularis oculi muscle, a muscle around the eye * Orbicularis oris muscle, a muscle around the mouth * Orbicular batfish, a specie ...
to ovate and is often held roughly horizontally, at a right angle to the pitcher orifice. It has a rounded to
emarginate 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 o ...
apex and a slightly
cordate Cordate is an adjective meaning 'heart-shaped' and is most typically used for: * Cordate (leaf shape), in plants * Cordate axe, a prehistoric stone tool See also * Chordate A chordate () is an animal of the phylum Chordata (). All chordat ...
base, and measures up to 4 cm in length by 3 cm in width. It has no appendages, but bears numerous nectar glands, which are scattered quite evenly across the entire lower surface of the lid. These nectaries are circular to shortly elliptic and thinly bordered, measuring around 0.3 mm in diameter. They become slightly larger and more densely packed around the midline. Their rims are visibly asymmetric and are highest near the apical end of the lid. Three to four prominent veins are present on either side of the lid's midline. The spur is broad and flattened, measuring up to 10 mm in length. It has 2–4 acute points at its apex, and has been variously described as either simple or branched.


Inflorescence

''Nepenthes aristolochioides'' 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 stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
up to 30 cm long. Both the peduncle and
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 ...
may be up to 15 cm long, although the latter is usually shorter in female plants. The peduncle is up to 4 mm in diameter.
Pedicel Pedicle or pedicel may refer to: Human anatomy *Pedicle of vertebral arch, the segment between the transverse process and the vertebral body, and is often used as a radiographic marker and entry point in vertebroplasty and kyphoplasty procedures ...
s are simple-
bract In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...
eolate and one-flowered. The basalmost ones are up to 12 mm long, whereas those higher up the rachis reach only 6 mm.
Tepal A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...
s are ovate and up to 4 mm long.
Fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
s are up to 20 mm long and 4 mm wide, and bear lanceolate valves.
Seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
s are
filiform Filiform, thread or filament like, can refer to: *Filiform, a common term used in botany to describe a thread-like shape *Filiform, or filiform catheter, a medical device whose component parts or segments are all cylindrical and more or less uni ...
.


Indumentum

Most parts of the plant are
glabrous Glabrousness (from the Latin ''glaber'' meaning "bald", "hairless", "shaved", "smooth") is the technical term for a lack of hair, down, setae, trichomes or other such covering. A glabrous surface may be a natural characteristic of all or part of ...
. Where present, the
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 ...
is inconspicuous; hairs are found on the
leaf axil A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
s, midribs, laminar margins, and parts of the pitchers (especially around the peristome and on the lid, and in developing pitchers). The indumentum is sparse and consists of short, simple or irregularly branching, appressed hairs, which are white to silver in colour and measure up to 0.2 mm and sometimes even 2 mm in length.


Colouration

The stem, laminae, tendrils and midribs are yellowish-green. On their outer surfaces, pitchers are white to reddish with numerous reddish-brown to purple speckles, with both lower and upper pitchers exhibiting similar colouration. The dark blotches are often denser in the upper part of the pitcher, though the extent of the translucent lighter patches is almost twice as great on the rear of the pitcher as compared to the front. The peristome is usually dark red or purple, being particularly dark in rosette pitchers. The undersurface of the lid is dark red or purple throughout, while the upper surface is speckled like the rest of the pitcher. The inner surface of the pitcher is white to light yellow throughout. Herbarium specimens are brown to dark brown, the preserved pitchers having conspicuous dark spots. Little variation has been observed within natural populations of ''N. aristolochioides'' and no
infraspecific taxa In botany, an infraspecific name is the scientific name for any taxon below the rank of species, i.e. an infraspecific taxon or infraspecies. (A "taxon", plural "taxa", is a group of organisms to be given a particular name.) The scientific names ...
have been described.


Ecology


Distribution and habitat

''Nepenthes aristolochioides'' is
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 elsew ...
to
Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
and has an altitudinal distribution of 1800–2500 m above sea level.McPherson, S.R. & A. Robinson 2012. ''
Field Guide to the Pitcher Plants of Sumatra and Java 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 taxonomic monographs released as part of larger ...
''. Redfern Natural History Productions, Poole.
In 2001,
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 Th ...
wrote that the species was only known from
Mount Tujuh Mount Tujuh ( id, Gunung Tujuh) is a caldera volcano in the Barisan Mountains of Sumatra. It has seven (tujuh) peaks, of which only three have been climbed.Clarke, C.M. 2001. ''Nepenthes of Sumatra and Peninsular Malaysia''. Natural History Public ...
in
Jambi Jambi is a province of Indonesia. It is located on the east coast of central Sumatra and spans to the Barisan Mountains in the west. Its capital and largest city is Jambi. The province has a land area of 50,160.05 km2, and a sea area of 3, ...
, although specimens collected by
Herbert Christopher Robinson Herbert Christopher Robinson (4 November 1874 – 20 May 1929) was a British zoologist and ornithologist. He is principally known for conceiving and initiating the major ornithological reference '' The Birds of the Malay Peninsula''. Robinson ...
and
Cecil Boden Kloss Cecil Boden Kloss (28 March 1877 – 19 August 1949) was an English zoologist. He was an expert on the mammals and birds of Southeast Asia. The Rubiaceae genus '' Klossia'' was named after him. Kloss was born in a family of Dutch descent who liv ...
labelled as being taken from "Mt. Kerinci" suggested that it may be more widespread in the region.
Mount Kerinci Mount Kerinci (also spelled Kerintji, among several other ways, and referred to as Gunung Kerinci, Gadang, Berapi Kurinci, Kerinchi, Korinci/Korintji, or Peak of Indrapura/Indrapoera) is an active stratovolcano and the highest mountain in Sumatra ...
is Sumatra's highest peak and neighbours Mount Tujuh. Clarke noted that since most of the mountain remains unexplored, there is a good chance that ''N. aristolochioides'' occurs there as well. The full range of ''N. aristolochioides'' on Mount Tujuh is also unknown, since only three of the mountain's seven peaks had been climbed as of 2001. In 2009, Stewart McPherson reported that ''N. aristolochioides'' was known from three subpopulations in
Kerinci Seblat National Park Kerinci Seblat National Park is the largest national park on the island of Sumatra, Indonesia. It has a total area of 13,791 km2, and spans four provinces: West Sumatra, Jambi, Bengkulu, and South Sumatra. Geography It is located between 1 ...
.McPherson, S. 2009. ''Nepenthes aristolochioides'' and ''Nepenthes klossii''. Two of Indonesia's most remarkable plants. ''
Carniflora Australis ''Carniflora Australis'' was a biannual English-language periodical and the official publication of the Australasian Carnivorous Plant Society.Rice, B. 2010Carnivorous Plant Society Archives The Carnivorous Plant FAQ. rchived page from October 1 ...
'' 7(1): 27–30.
Of these, one (the type locality on Mount Tujuh) may already be
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
due to poaching by
plant collector Plant collecting is the acquisition of plant specimens for the purposes of research, cultivation, or as a hobby. Plant specimens may be kept alive, but are more commonly dried and pressed to preserve the quality of the specimen. Plant collecting i ...
s; in 2007, E'En Endatno observed only a single ''N. aristolochioides'' plant on Mount Tujuh. The other two sites are located on remote peaks and support only "a few dozen" plants, as determined by McPherson. ''Nepenthes aristolochioides'' inhabits ''
Sphagnum ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, peat moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store wa ...
''-dominated
mossy forest A cloud forest, also called a water forest, primas forest, or tropical montane cloud forest (TMCF), is a generally tropical or subtropical, evergreen, montane, moist forest characterized by a persistent, frequent or seasonal low-level cloud c ...
near the tops of steep ridges. It usually grows terrestrially, but can also occur as an
epiphyte An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
in pockets of moss on tree trunks. The species occurs sympatrically with '' N. gymnamphora'' and '' N. singalana''. It grows with the former in
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 f ...
and
swamp A swamp is a forested wetland.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p. Swamps are considered to be transition zones because both land and water play a role in ...
s dominated by ''
Pandanus ''Pandanus'' is a genus of monocots with some 750 accepted species. They are palm-like, dioecious trees and shrubs native to the Old World tropics and subtropics. The greatest number of species are found in Madagascar and Malaysia. Common names ...
'' species that line the shoreline of a crater lake. The altitudinal distribution of ''N. gymnamphora'' on Mount Tujuh (1800–2100 m) overlaps that of ''N. aristolochioides'', but no
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 ...
s have been observed. A small form of ''N. singalana'' occurs in the same habitat as ''N. aristolochioides'', but appears to occupy a different
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
; it is generally confined to the forest floor while ''N. aristolochioides'' often climbs into the canopy. A number of plants representing the
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 ...
''N. aristolochioides'' × ''N. singalana'' have been recorded.


Conservation

''Nepenthes aristolochioides'' is listed as Critically Endangered on the 2013 IUCN Red List of Threatened Species, as its known distribution was restricted to two mountains (Mount Kerinci and Mount Tujuh) at the time of the assessment. Despite the fact that all known populations of the species lie within
Kerinci Seblat National Park Kerinci Seblat National Park is the largest national park on the island of Sumatra, Indonesia. It has a total area of 13,791 km2, and spans four provinces: West Sumatra, Jambi, Bengkulu, and South Sumatra. Geography It is located between 1 ...
, it is severely threatened by over-collection, because its unique pitcher morphology makes it particularly sought-after. In 2010, the Rare ''Nepenthes'' Collection was established with the aim of conserving 4 of the most endangered ''Nepenthes'' species: ''N. aristolochioides'', '' N. clipeata'', '' N. khasiana'', and '' N. rigidifolia''.Bourke, G. 2010
A new conservation initiative: the Rare ''Nepenthes'' Collection project.
''Captive Exotics Newsletter'' 1(2): 5–6.


Pitcher infauna

No infaunal organisms have been recorded from the pitchers of ''N. aristolochioides''. This is not due to a lack of potential inhabitants; pitchers of '' N. singalana'', which grow alongside ''N. aristolochioides'', support large populations of such organisms. It is thought that the structure of the traps may serve to disorientate emerging adults and so infaunal species avoid colonising them.


Carnivory

Different trapping mechanisms have been proposed for the lower and upper pitchers of ''N. aristolochioides''.


Pitfall traps

The lower pitchers of this species frequently develop embedded in ''
Sphagnum ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, peat moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store wa ...
'' moss, with only the top of the traps visible.
Joachim Nerz Dr. Joachim Nerz (born 1964) is a German taxonomist and botanist specialising in the carnivorous plant genera ''Heliamphora'' and '' Nepenthes''. Nerz has described several new species, mostly with Andreas Wistuba. Publications * Schlauer, J. ...
suggested that they act as simple
pitfall traps A pitfall trap is a trapping pit for small animals, such as insects, amphibians and reptiles. Pitfall traps are a sampling technique, mainly used for ecology studies and ecologic pest control. Animals that enter a pitfall trap are unable to esca ...
specialised for trapping ground-dwelling insects. The insects crawl into the pitcher through the small mouth and fall to the bottom of the pitcher cup. Unable to climb out, they drown in the digestive fluid.


Light traps

Along with '' N. klossii'', ''N. aristolochioides'' is the only species in the genus to employ domed pitchers with translucent patches that allow sunlight to illuminate the interior. When viewed from the front, the peristome and lid appear dark, contrasting heavily against the inner surface of the pitcher, which is brightly lit by light passing through the top of the pitcher dome. It has been suggested that in upper pitchers this adaptation serves to attract flying insects in a similar manner to the
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
n pitcher plants ''
Darlingtonia californica ''Darlingtonia californica'' , also called the California pitcher plant, cobra lily, or cobra plant, is a species of carnivorous plant. It is the sole member of the genus ''Darlingtonia'' in the family Sarraceniaceae. This pitcher plant is nativ ...
'', ''
Sarracenia minor ''Sarracenia minor'', also known as the hooded pitcherplant, is a perennial, terrestrial, rhizomatous, herbaceous, carnivorous plant in the genus ''Sarracenia''. Like all the ''Sarracenia'', it is native to North America. Etymology In 1788, the ...
'', and ''
Sarracenia psittacina ''Sarracenia psittacina'', also known as the parrot pitcherplant, is a carnivorous plant in the genus ''Sarracenia''. Like all the ''Sarracenia'', it is native to North America, in the Southeastern United States. ''Sarracenia psittacina'' emplo ...
''. A similar trapping mechanism has also been proposed for '' N. jacquelineae''. This is supported by the fact that most of the prey caught by ''N. aristolochioides'' consist of small
flies Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
, which are attracted to bright light sources. Prey are often disorientated inside the upper pitchers of ''N. aristolochioides''. Unable to find the exit, they eventually fall into the pitcher fluid and drown. As such, the pitchers have features of lobster-pot traps. The central role of the translucent dome in the prey trapping mechanism of ''N. aristolochioides'' is supported by experimental evidence. In one study, pitchers whose domes were covered with red celluloid filters showed a threefold decrease in ''
Drosophila ''Drosophila'' () is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or (less frequently) pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species ...
'' trapping efficiency as compared to unaltered pitchers and those shaded at the front with the same filters (flies are red-blind and most sensitive to the UV, blue, and green wavebands).Moran, J.A., C. Clarke & B.E. Gowen 2012. The use of light in prey capture by the tropical pitcher plant ''Nepenthes aristolochioides''. ''Plant Signaling & Behavior'' 7(8): 957–960.


Flypaper traps

''Nepenthes aristolochioides'' produces extremely thick,
mucilaginous Mucilage is a thick, gluey substance produced by nearly all plants and some microorganisms. These microorganisms include protists which use it for their locomotion. The direction of their movement is always opposite to that of the secretion of m ...
pitcher liquid, which coats the entire inner surfaces of the traps in a thin film. The pitchers of this species appear to function at least in part as flypaper traps, with the sticky inner walls trapping flying insects above the surface of the fluid. Similarly viscous pitcher fluid is also found in seven other closely allied
Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
n species: '' N. dubia'', '' N. flava'', '' N. inermis'', '' N. jacquelineae'', '' N. jamban'', '' N. talangensis'', and '' N. tenuis''. Together with ''N. aristolochioides'', these species all share
infundibular An infundibulum (Latin for ''funnel''; plural, ''infundibula'') is a funnel-shaped cavity or organ. Anatomy * Brain: the pituitary stalk, also known as the ''infundibulum'' and ''infundibular stalk'', is the connection between the hypothalamus and ...
pitchers that are wholly glandular or almost so.


Related species

The unusual pitcher morphology of ''N. aristolochioides'' makes it difficult to confuse with any other species; the almost vertical orientation of the pitcher mouth is a unique characteristic.
Joachim Nerz Dr. Joachim Nerz (born 1964) is a German taxonomist and botanist specialising in the carnivorous plant genera ''Heliamphora'' and '' Nepenthes''. Nerz has described several new species, mostly with Andreas Wistuba. Publications * Schlauer, J. ...
noted that ''N. aristolochioides'' shows "close affinities" to '' N. talangensis''. However, it may be easily distinguished from that species on the basis of the pitcher mouth, which is horizontal in ''N. talangensis''. In addition, the pitcher mouth of ''N. talangensis'' is elongated into a short neck, whereas ''N. aristolochioides'' lacks a neck altogether, with the lid being inserted in front of the pitcher. Both the mouth and lid are considerably larger in ''N. talangensis''. The two
taxa 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 nam ...
also differ somewhat in growth habit; ''N. talangensis'' occurs only terrestrially and is a weak climber, whereas ''N. aristolochioides'' occasionally grows as an
epiphyte An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
and climbs high into the forest canopy.Nerz, J. & A. Wistuba 1994
Five new taxa of ''Nepenthes'' (Nepenthaceae) from North and West Sumatra
. ''
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 ...
'' 23(4): 101–114.
The laminar morphology of ''N. aristolochioides'' is also similar to that of '' N. bongso'', although ''N. aristolochioides'' is easily distinguished from this species by the shape of its pitchers and the hooded nectaries of the lid. In 2001,
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 Th ...
performed a
cladistic analysis Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived chara ...
of the ''Nepenthes'' species of Sumatra and
Peninsular Malaysia Peninsular Malaysia ( ms, Semenanjung Malaysia; Jawi: سمننجڠ مليسيا), or the States of Malaya ( ms, Negeri-negeri Tanah Melayu; Jawi: نڬري-نڬري تانه ملايو), also known as West Malaysia or the Malaysian Peninsula, ...
using 70 morphological characteristics of each taxon. The following is part of the resultant
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
, showing "Clade 1", which has 51% bootstrap support. Its most strongly supported subclade is the sister pair of ''N. inermis'' and ''N. dubia'', having 95% support. Although ''N. aristolochioides'' resembles '' N. klossii'' in some respects, the two species are geographically isolated from each other and are not thought to be closely related. The unique adaptations of these taxa might represent an example of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
, whereby two organisms that are not closely related independently acquire similar characteristics while evolving in separate, but comparable, ecosystems.


Natural hybrids

Only one natural
hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two dif ...
involving ''N. aristolochioides'' is known. ''Nepenthes aristolochioides'' × '' N. singalana'' has been found in dense
mossy forest A cloud forest, also called a water forest, primas forest, or tropical montane cloud forest (TMCF), is a generally tropical or subtropical, evergreen, montane, moist forest characterized by a persistent, frequent or seasonal low-level cloud c ...
on two ridges of
Mount Tujuh Mount Tujuh ( id, Gunung Tujuh) is a caldera volcano in the Barisan Mountains of Sumatra. It has seven (tujuh) peaks, of which only three have been climbed.Clarke, C.M. 2001. ''Nepenthes of Sumatra and Peninsular Malaysia''. Natural History Public ...
, only one of which is populated by ''N. aristolochioides''. It is relatively rare, which suggests that the two species flower at different times of the year. This hybrid is smaller than either of its parent species; the pitchers rarely exceed 5 cm in height. The lower pitchers resemble those of '' N. talangensis'', but differ in having more pronounced peristome teeth. Upper pitchers are infundibular in the lower parts, ovoid in the middle, and cylindrical in the upper parts. This hybrid can be distinguished from ''N. aristolochioides'' on the basis of its narrow, cylindrical peristome and oblique mouth, as opposed to almost vertical in the latter.


References


Further reading

* Clarke, C. 2013
What Can Tree Shrews Tell Us about the Effects of Climate Change on Pitcher Plants?
ideo IDEO () is a design and consulting firm with offices in the U.S., England, Germany, Japan, and China. It was founded in Palo Alto, California, in 1991. The company's 700 staff uses a design thinking approach to design products, services, environ ...
TESS seminars, 25 September 2013. * Harwood, P. 1998. Four ''Nepenthes'' "new to science". ''
The Carnivorous Plant Society Journal ''Planta Carnivora'' is a biannual periodical and the official publication of The Carnivorous Plant Society of the United Kingdom.Rice, B. 2010Carnivorous Plant Society Archives The Carnivorous Plant FAQ. rchived page from October 10, 2010/ref>< ...
'' 21: 64–65. * Hernawati & P. Akhriadi 2006. ''
A Field Guide to the Nepenthes of Sumatra 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 taxonomic monographs released as part of larger ...
''. PILI-NGO Movement, Bogor. * Meimberg, H., A. Wistuba, P. Dittrich & G. Heubl 2001. Molecular phylogeny of Nepenthaceae based on cladistic analysis of plastid trnK intron sequence data. ''Plant Biology'' 3(2): 164–175. * Meimberg, H. 2002.
Molekular-systematische Untersuchungen an den Familien Nepenthaceae und Ancistrocladaceae sowie verwandter Taxa aus der Unterklasse Caryophyllidae s. l..
Ph.D. thesis, Ludwig Maximilian University of Munich, Munich. * Meimberg, H. & G. Heubl 2006. Introduction of a nuclear marker for phylogenetic analysis of Nepenthaceae. ''Plant Biology'' 8(6): 831–840. * Meimberg, H., S. Thalhammer, A. Brachmann & G. Heubl 2006. Comparative analysis of a translocated copy of the ''trnK'' intron in carnivorous family Nepenthaceae. ''Molecular Phylogenetics and Evolution'' 39(2): 478–490.


External links


Photographs of ''N. aristolochioides''
at the Carnivorous Plant Photofinder {{Taxonbar, from=Q139559 Carnivorous plants of Asia aristolochioides Endemic flora of Sumatra Critically endangered flora of Asia Species endangered by the pet trade Taxa named by Matthew Jebb Taxa named by Martin Cheek