Nepenthes leonardoi
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''Nepenthes leonardoi'' 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 from a single locality in central
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 ...
, 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 ...
. It is closely allied to several other Palawan
endemics 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 ...
, including '' N. deaniana'', '' N. gantungensis'', and '' N. mira''. The traps of this species reach at least 24 cm in height. Some specimens are noted for producing very dark, almost black, upper pitchers. 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. leonardoi'' as a
heterotypic synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linnae ...
of '' N. deaniana''.


Botanical history

''Nepenthes leonardoi'' was discovered on November 18, 2010, by Greg Bourke, Jehson Cervancia, Mark Jaunzems, and Stewart McPherson. The species was initially known under the
placeholder name Placeholder names are words that can refer to things or people whose names do not exist, are tip of the tongue, temporarily forgotten, are not relevant to the salient point at hand, are to avoid stigmatization, are unknowable/unpredictable in ...
"''Nepenthes'' sp. Palawan".New Species - ''Nepenthes'' sp. "Palawan"
Redfern Natural History Productions. rchived page from July 15, 2011/ref> ''Nepenthes leonardoi'' was formally described in the March 2011 issue of ''
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 ...
'' by McPherson, Bourke, Cervancia, Jaunzems, and
Alastair Robinson Alastair S. Robinson (born 1980) is a taxonomist and field botanist specialising in the carnivorous plant genus ''Nepenthes'', for which he is regarded as a world authority.Ellison, A. & Adamec, L. eds., 2017. Contributing Author Information ...
. 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 ...
''leonardoi'' honours Filipino botanist
Leonardo Co Leonardo Legaspi Co (December 29, 1953 in Manila – November 15, 2010 in Kananga, Leyte) was a Filipino botanist and plant taxonomist who was considered the "foremost authority in ethnobotany in the Philippines" during his lifetime. Education ...
, who was killed on the island of
Leyte Leyte ( ) is an island in the Visayas group of islands in the Philippines. It is eighth-largest and sixth-most populous island in the Philippines, with a total population of 2,626,970 as of 2020 census. Since the accessibility of land has be ...
on November 15, 2010, reportedly as a result of "crossfire between government security forces and an insurgent group".Anda, R. 2010
New plant species named after Co
. ''Inquirer Southern Luzon'', December 24, 2010.
The date of Co's death coincided with the first day of the ''Nepenthes'' expedition that led to the discovery of ''N. leonardoi''. As explained in the species's describing paper:
The first four authors of this species discovered this plant on November 18th, and felt it fitting to name this plant, which is unique among Philippine ''Nepenthes'' in producing black pitchers, after Leonardo, in honour of his lifework and many accomplishments.
The
herbarium A herbarium (plural: herbaria) is a collection of preserved plant specimens and associated data used for scientific study. The specimens may be whole plants or plant parts; these will usually be in dried form mounted on a sheet of paper (called ...
specimen ''S. McPherson SRM 5'' is the designated
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 ...
, and is deposited at the herbarium of
Palawan State University The Palawan State University also referred to by its acronym PSU is a public government-funded higher education institution in Puerto Princesa, Palawan, Philippines. It was established History The Palawan State University, the first state unive ...
(PPC),
Puerto Princesa City Puerto Princesa, officially the City of Puerto Princesa ( Cuyonon: ''Siyudad i'ang Puerto Princesa''; fil, Lungsod ng Puerto Princesa), is a 1st class highly urbanized city in the Mimaropa region of the Philippines. According to the 2020 censu ...
. It was collected on November 20, 2010, near the summit of Schom-carp Peak at 1490 m. In preparing the species description, the describing authors also examined herbarium material of a number of closely allied species, including '' N. attenboroughii'' (specimens ''A.Robinson AR001'' and ''AR002''), '' N. mantalingajanensis'' (''G.C.G.Argent & E.M.Romero 92114''), '' N. mira'' (''G.C.G.Argent et al. 25438''), and '' N. rajah'' (''Low s.n.'').


Description

''Nepenthes leonardoi'' is a climbing or scrambling plant. The
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
, which is unbranched, reaches a maximum length of around 4 m. It is cylindrical and varies in diameter from 1.5–2.8 cm. Internodes are typically 1.5–18 cm long, becoming elongated in climbing specimens. Plants found under the shade of dense vegetation typically have longer internodes compared to those growing in more open areas, although the latter on average bear larger pitchers and
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 ...
s.


Leaves

Leaves are
petiolate Petiole may refer to: *Petiole (botany), the stalk of a leaf, attaching the blade to the stem *Petiole (insect anatomy) In entomology, petiole is the technical term for the narrow waist of some hymenopteran insects, especially ants, bees, a ...
to subpetiolate and
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 ...
in texture. In mature plants, 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 ...
(leaf blade) is narrowly oblong and measures 15–50 cm in length by 6–10 cm in width. The laminar apex is typically acute or rounded, but may occasionally be abruptly truncated. The base may be shortly attenuate or obtuse, and encircles the stem for two-thirds to the entirety of its circumference. In younger plants, the lamina is usually narrower towards the base, becoming more oblong as the plant matures. The apex may sometimes be slightly
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 ...
, with the tendril joining the lamina on the underside, up to 4 mm before the apex. Occasionally the two halves of the lamina may meet 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 ...
unequally at different points along its length, up to 3 mm apart in some specimens.
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 may be extremely long, sometimes exceeding 130 cm, particularly on leaves bearing lower pitchers. Those supporting upper pitchers are coiled.


Pitchers

Rosette and lower pitchers are variable in shape: they are usually wholly ovate or
urceolate 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 may also be
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 ...
. They grow up to 15 cm high by 6 cm wide, but are often considerably smaller. A pair of wings up to 12 mm wide runs down 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 ...
surface of the pitcher cup. These wings bear
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 ...
fringe elements up to 10 mm long. The pitcher mouth is oval to circular and up to 6 cm across. It rises at the rear to form a conspicuous neck. 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 mosses, ...
is cylindrical to slightly flattened and up to 2 cm wide for most of its length, becoming broader towards the neck (≤2.5 cm wide). It bears ribs up to 2 mm high and spaced up to 2 mm apart, which terminate in teeth (≤4 mm long) on the peristome's inner margin. The digestive zone of the inner surface usually extends for around two-thirds of the pitcher's height, although some specimens may be wholly glandular. The pitcher lid or operculum is elliptic and measures up to 5.5 cm in length by 3.5 cm in width. It lacks appendages or a keel. A
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 ...
is inserted near the base of the lid. It measures up to 9 mm in length, with a basal diameter of up to 3 mm, although it may be much smaller. Upper pitchers are variably funnel-shaped, ranging from wholly
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 ...
to infundibular only in the basal quarter and cylindrical, or more rarely tubular, above. They may be considerably larger than their terrestrial counterparts, reaching 24 cm in height by 6 cm in width, though they rarely approach these maximum dimensions. The wings are commonly reduced to a pair of ribs in aerial traps. Upper pitchers resemble lower ones in other respects.


Inflorescence

''Nepenthes leonardoi'' is known to flower both in the rosette stage and as a vining plant. The species 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 ...
. Male inflorescences can reach a height of 50 cm, of which 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 ...
constitutes up to 30 cm. Female inflorescences are similar in size, typically growing to 45 cm in length. Generally both male and female inflorescences are much shorter, however, especially in the case of plants growing in exposed sites. This being the case, exceptional specimens (also found in exposed areas) may produce a rigid inflorescence up to 110 cm length, of which only the distal 15% bears flowers. Inflorescences of both sexes have a basal diameter of approximately 1 cm and hold up to around 120 closely packed flowers, which are usually restricted to the distal quarter to half of its length. Flowers are usually borne on one-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 ...
, although two-flowered partial peduncles may also be present. The inflorescence often bears a vestigial leaf below the rachis. Male flowers are fragrant; their scent has been described as "distinctive, musty, sweet" and is noticeable from a distance of up to 60 cm. The smell of female flowers has not been recorded.
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 8 mm long. As in most ''Nepenthes'' species, the seeds are filiform. They are pale brown and around 7 mm long.


Indumentum

A sparse
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 simple, reddish-brown hairs is present on certain vegetative parts, including the outside of the pitchers, the laminar margins, the lower surface of the midrib, and the tendrils (where they are somewhat more densely distributed). These hairs may be 2 mm long, but are typically shorter. They are
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 ...
, being lost with age. Consequently, the laminar margins and midrib are particularly
hirsute Hirsutism is excessive body hair on parts of the body where hair is normally absent or minimal. The word is from early 17th century: from Latin ''hirsutus'' meaning "hairy". It usually refers to a "male" pattern of hair growth in a female that ...
in developing leaves.


Colour

Colouration is highly variable. The laminae and petioles are often green throughout when produced in shaded conditions and red to purple in plants exposed to direct sunlight. In certain specimens the underside of the lamina and/or the stem may be red to purple, but this is uncommon. A red tinge to the stem is already apparent in many juvenile plants. The range of colours displayed by the pitchers of ''N. leonardoi'' is exceptional among Philippine ''Nepenthes''. Lower pitchers are most commonly orange to red on their outer surfaces, often flecked with slightly darker purple markings. The wings may match the external pigmentation of the pitcher cup or be yellowish-green. The peristome is often red and may turn purple with age. The lid is yellow to orange, frequently with red highlights. In aerial traps, the pitcher cup is predominantly yellowish-green with a darker, orange to red peristome. In certain specimens the upper pitchers may be faintly red speckled. Beyond this common pigmentation, both lower and upper pitchers exhibit a continuum of colours across the species, from wholly yellowish-green at one extreme to entirely
burgundy Burgundy (; french: link=no, Bourgogne ) is a historical territory and former administrative region and province of east-central France. The province was once home to the Dukes of Burgundy from the early 11th until the late 15th century. The c ...
at the other. The latter colour form is particularly striking and is far more frequently encountered in ''N. leonardoi'' than in any other closely allied Philippine ''Nepenthes''. In extreme cases, the upper pitchers may look almost completely black. Flash photography shows the actual colour of the pitcher cup to be a uniform deep purple; their very dark appearance seems to result from a combination of this purple pigmentation and their brown indumentum. This effect is particularly pronounced in wet pitchers.


Ecology

''Nepenthes leonardoi'' 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 the
Philippine 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 ...
island of
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 ...
. It has only been recorded from the upper slopes of a single mountain system centred on Schom-carp Peak (also known as ''Shumkat Peak'' or ''Shumkak Peak''); it grows on the peak itself as well as on its eastern ridge, which extends in the direction of Narra
municipality A municipality is usually a single administrative division having corporate status and powers of self-government or jurisdiction as granted by national and regional laws to which it is subordinate. The term ''municipality'' may also mean the go ...
. The presence of this species on the easterly ridge connecting Schom-carp Peak with the Brow Shoulder
massif In geology, a massif ( or ) is a section of a planet's crust that is demarcated by faults or flexures. In the movement of the crust, a massif tends to retain its internal structure while being displaced as a whole. The term also refers to a ...
could not be confirmed by the describing authors, but they suggested it is likely to grow there and possibly on the massif itself. The altitudinal distribution of ''N. leonardoi'' ranges from at least 1300 m
above sea level Height above mean sea level is a measure of the vertical distance (height, elevation or altitude) of a location in reference to a historic mean sea level taken as a vertical datum. In geodesy, it is formalized as ''orthometric heights''. The comb ...
to the summit of Schom-carp at 1490 m. The only known population numbers several thousand individuals. ''Nepenthes leonardoi'' is exclusively
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
. Its typical habitat is upper
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 ...
(on the eastern ridge) and upper montane scrub (in the summit area of Schom-carp Peak). It is not
sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species sh ...
with other ''Nepenthes'' species and 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 recorded. Like its close relatives, ''N. leonardoi'' is apparently outcompeted by
bamboo Bamboos are a diverse group of evergreen perennial flowering plants making up the subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family. The origin of the word "bamboo" is uncertain, bu ...
and is consequently largely or completely absent from the highest parts of the eastern ridge. The infaunal community of ''N. leonardoi'' pitchers includes small
spider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
sThe Discovery of ''Nepenthes leonardoi''
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Redfern Natural History Productions.
and unidentified
mosquito Mosquitoes (or mosquitos) are members of a group of almost 3,600 species of small flies within the family Culicidae (from the Latin ''culex'' meaning " gnat"). The word "mosquito" (formed by ''mosca'' and diminutive ''-ito'') is Spanish for "li ...
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e, the latter being found in abundance in both lower and upper pitchers. The
conservation status The conservation status of a group of organisms (for instance, a species) indicates whether the group still exists and how likely the group is to become extinct in the near future. Many factors are taken into account when assessing conservation ...
of ''N. leonardoi'' has not been formally assessed for the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
, but the describing authors suggested it be considered Critically Endangered based on the IUCN criteria, due to its localised distribution and small population size. Although the habitat of ''N. leonardoi'' remains untouched by human activity, the describing authors cited mining operations to the south, on the Mount Victoria massif (to which '' N. attenboroughii'' is endemic), as a possible future threat if appropriate conservation measures are not taken.


Notes

:a.Similar outcomes of
interspecific competition Interspecific competition, in ecology, is a form of competition in which individuals of ''different'' species compete for the same resources in an ecosystem (e.g. food or living space). This can be contrasted with mutualism, a type of symbiosis. ...
with bamboo have been observed in '' N. attenboroughii'', '' N. deaniana'', '' N. gantungensis'', '' N. mantalingajanensis'', '' N. mira'', and '' N. palawanensis''.


References


Further reading

* Co, L. & W. Suarez 2012
Nepenthaceae
Co's Digital Flora of the Philippines. * McPherson, S.R. & V.B. Amoroso 2011. ''
Field Guide to the Pitcher Plants of the Philippines 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 larg ...
''. Redfern Natural History Productions, Poole. * McPherson, S.R. 2011. Comparison of the highland Palaweño ''Nepenthes''. In: '' New Nepenthes: Volume One''. Redfern Natural History Productions, Poole. pp. 364–381. * McPherson, S.R. 2012. ''Nepenthes leonardoi''. In: Associazione Italiana Piante Carnivore. p. 21.
Exploration of Mount Anipahan and Mount Kiamo
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 ...
Redfern Natural History Productions. {{Taxonbar, from=Q6994724 Carnivorous plants of Asia leonardoi Endemic flora of the Philippines Flora of Palawan Critically endangered flora of Asia Plants described in 2011