Heliamphora
Encyclopedia
The genus Heliamphora contains 23 species of pitcher plant
Pitcher plant
Pitcher plants are carnivorous plants whose prey-trapping mechanism features a deep cavity filled with liquid known as a pitfall trap. It has been widely assumed that the various sorts of pitfall trap evolved from rolled leaves, with selection pressure favouring more deeply cupped leaves over...

s endemic to South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...

. The species are collectively known as sun pitchers, based on the mistaken notion that the heli of Heliamphora is from the Greek helios, meaning "sun". In fact, the name derives from helos, meaning marsh, so a more accurate translation of their scientific name would be marsh pitcher plants. Species in the genus Heliamphora are carnivorous plant
Carnivorous plant
Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans, typically insects and other arthropods. Carnivorous plants appear adapted to grow in places where the soil is thin or poor in nutrients, especially nitrogen, such as acidic...

s that consist of a modified leaf form that is fused into a tubular shape. They have evolved mechanisms to avoid completely filling up with water and attract, trap, and kill insects. At least one species (H. tatei
Heliamphora tatei
Heliamphora tatei is a species of Marsh Pitcher Plant endemic to Duida Tepui, Cerro Huachamacari and Marahuaca Tepui in Venezuela. It is closely related to H. neblinae and the latter was at one point considered a variety of H. tatei.-Infraspecific taxa:*Heliamphora tatei var. macdonaldae Maguire ...

) produces its own proteolytic
Proteolysis
Proteolysis is the directed degradation of proteins by cellular enzymes called proteases or by intramolecular digestion.-Purposes:Proteolysis is used by the cell for several purposes...

 enzymes that allows it to digest its prey without the help of symbiotic bacteria.

Morphology

All Heliamphora are herbaceous
Herbaceous
A herbaceous plant is a plant that has leaves and stems that die down at the end of the growing season to the soil level. They have no persistent woody stem above ground...

 perennial plant
Perennial plant
A perennial plant or simply perennial is a plant that lives for more than two years. The term is often used to differentiate a plant from shorter lived annuals and biennials. The term is sometimes misused by commercial gardeners or horticulturalists to describe only herbaceous perennials...

s that grow from a subterranean rhizome
Rhizome
In botany and dendrology, a rhizome is a characteristically horizontal stem of a plant that is usually found underground, often sending out roots and shoots from its nodes...

. H. tatei grows as a shrub
Shrub
A shrub or bush is distinguished from a tree by its multiple stems and shorter height, usually under 5–6 m tall. A large number of plants may become either shrubs or trees, depending on the growing conditions they experience...

, up to four meters tall, all other species form prostrate rosettes. The leaf size ranges from a few centimeters (H. minor
Heliamphora minor
Heliamphora minor is a species of Marsh Pitcher Plant endemic to the Auyan and Chimanta group of tepuis in Venezuela. As the name suggests, it is the smallest species in the genus. It is closely related to H. pulchella....

, H. pulchella
Heliamphora pulchella
Heliamphora pulchella is a species of Marsh Pitcher Plant endemic to the Chimanta group of tepuis in Venezuela. It is one of the smallest species and closely related to H...

) up to more than 50 cm (H. ionasi). Heliamphora possess tubular traps formed by rolled leaves with fused edges. Marsh pitcher plants are unusual among pitcher plant
Pitcher plant
Pitcher plants are carnivorous plants whose prey-trapping mechanism features a deep cavity filled with liquid known as a pitfall trap. It has been widely assumed that the various sorts of pitfall trap evolved from rolled leaves, with selection pressure favouring more deeply cupped leaves over...

s in that they lack lids (opercula
Operculum (botany)
An operculum, in botany, is a term generally used to describe a structure within a plant, moss, or fungus acting as a cap, flap, or lid. In plants, it may also be called a bud cap.Examples of structures identified as opercula include:...

), instead having a small "nectar spoon" on the upper posterior portion of the leaf. This spoon-like structure secretes a nectar-like substance, which serves as a lure for insects and small animals. Each pitcher also exhibits a small slit in its side that allows excess rainwater to drain away, similar to the overflow on a sink. This allows the marsh pitcher plants to maintain a constant maximum level of rainwater within the pitcher. The pitchers' inner surface is covered with downward-pointing hairs to force insects into the pitchers' lower parts.

Carnivory

Though often counted among the various carnivorous plant
Carnivorous plant
Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans, typically insects and other arthropods. Carnivorous plants appear adapted to grow in places where the soil is thin or poor in nutrients, especially nitrogen, such as acidic...

s, with the exception of Heliamphora tatei, the vast majority of plants in the genus Heliamphora do not produce their own digestive enzymes (i.e. protease
Protease
A protease is any enzyme that conducts proteolysis, that is, begins protein catabolism by hydrolysis of the peptide bonds that link amino acids together in the polypeptide chain forming the protein....

s, ribonuclease
Ribonuclease
Ribonuclease is a type of nuclease that catalyzes the degradation of RNA into smaller components. Ribonucleases can be divided into endoribonucleases and exoribonucleases, and comprise several sub-classes within the EC 2.7 and 3.1 classes of enzymes.-Function:All organisms studied contain...

s, phosphatase
Phosphatase
A phosphatase is an enzyme that removes a phosphate group from its substrate by hydrolysing phosphoric acid monoesters into a phosphate ion and a molecule with a free hydroxyl group . This action is directly opposite to that of phosphorylases and kinases, which attach phosphate groups to their...

s, etc.), relying instead on the enzymes of symbiotic bacteria to break down their prey. They do, however, attract prey through special visual and chemical signals and trap and kill the prey through a typical pitfall trap. Field studies of H. nutans
Heliamphora nutans
Heliamphora nutans is a species of Marsh Pitcher Plant native to Venezuela, Brazil and Guyana, where it grows on several Tepuis, including Roraima, Kukenan, Tramen and Ilu. H...

, H. heterodoxa, H. minor, and H. ionasi have determined that none of these species produce their own proteolytic
Proteolysis
Proteolysis is the directed degradation of proteins by cellular enzymes called proteases or by intramolecular digestion.-Purposes:Proteolysis is used by the cell for several purposes...

 enzymes. If production of these enzymes was used as a strict demarcation of what is and what is not a carnivorous plant, many of the Heliamphora species would not meet the requirement. H. tatei is one of the few species observed to produce both digestive enzymes and wax scales, which also aid in prey capture. The pattern of carnivory among Heliamphora species, combined with habitat data, indicates that carnivory in this genus evolved in nutrient-poor locations as a means to improve absorption of available nutrients. Most Heliamphora typically capture ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...

s, while H. tateis improvements on the design allow it to capture and absorb nutrients from more flying insects. The carnivorous habit among these species is lost in low light conditions, which suggests that certain nutrient concentrations (specifically nitrogen
Nitrogen
Nitrogen is a chemical element that has the symbol N, atomic number of 7 and atomic mass 14.00674 u. Elemental nitrogen is a colorless, odorless, tasteless, and mostly inert diatomic gas at standard conditions, constituting 78.08% by volume of Earth's atmosphere...

 and phosphorus
Phosphorus
Phosphorus is the chemical element that has the symbol P and atomic number 15. A multivalent nonmetal of the nitrogen group, phosphorus as a mineral is almost always present in its maximally oxidized state, as inorganic phosphate rocks...

) are only limiting during periods of fast growth under normal light conditions, thus rendering most of the carnivorous adaptations inefficient and not energy cost effective.

Distribution

All Heliamphora species are endemic
Endemic (ecology)
Endemism is the ecological state of being unique to a defined geographic location, such as an island, nation or other defined zone, or habitat type; organisms that are indigenous to a place are not endemic to it if they are also found elsewhere. For example, all species of lemur are endemic to the...

 to the Tepui
Tepui
A tepui , or tepuy, is a table-top mountain or mesa found in the Guiana Highlands of South America, especially in Venezuela. The word tepui means "house of the gods" in the native tongue of the Pemon, the indigenous people who inhabit the Gran Sabana....

s of the Guiana Highlands in the tripoint Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

, Guyana
Guyana
Guyana , officially the Co-operative Republic of Guyana, previously the colony of British Guiana, is a sovereign state on the northern coast of South America that is culturally part of the Anglophone Caribbean. Guyana was a former colony of the Dutch and of the British...

 and Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...

, except H. heterodoxa
Heliamphora heterodoxa
Heliamphora heterodoxa is a species of Marsh Pitcher Plant native to Gran Sabana and the plateau of Ptari Tepui in Venezuela. It was first discovered in 1951 upon Ptari Tepui. This Heliamphora is closely related to H. glabra and the latter was for a long time considered a form of H...

 and H. sarracenioides
Heliamphora sarracenioides
Heliamphora sarracenioides is a species of Marsh Pitcher Plant native to southern Venezuela, Gran Sabana and Estado Bolivar.-References:...

, which also occur in the Gran Sabana. Many of the Tepuis have not yet been explored for Heliamphora, and the large number of species described in recent years suggests that many more species may be awaiting discovery.

Botanical history

The first species of the genus to be described was H. nutans, which George Bentham
George Bentham
George Bentham CMG FRS was an English botanist, characterized by Duane Isely as "the premier systematic botanist of the nineteenth century".- Formative years :...

 named in 1840 based on a specimen collected by Robert Hermann Schomburgk
Robert Hermann Schomburgk
Sir Robert Hermann Schomburgk , was a German-born explorer for Great Britain who carried out geographical, ethnological and botanical studies in South America and the West Indies, and also fulfilled diplomatic missions for Great Britain in the Dominican Republic and Thailand.-Biography:Schomburgk...

. This remained the only known species until Henry Allan Gleason
Henry Gleason
Henry Allan Gleason was a noted American ecologist, botanist, and taxonomist, most recognized for his endorsement of the individualistic/open community concept of ecological succession.- Life and work :...

 described H. tatei and H. tyleri in 1931, also adding H. minor in 1939. Between 1978 and 1984, Julian Alfred Steyermark
Julian Alfred Steyermark
Julian Alfred Steyermark was an American botanist. His focus was on New World vegetation, and he specialized in the family Rubiaceae.- Life and work :...

 and Bassett Maguire revised the genus (to which Steyermark had added H. heterodoxa in 1951) and described two more species, H. ionasi and H. neblinae
Heliamphora neblinae
Heliamphora neblinae is a species of Marsh Pitcher Plant endemic to Cerro Neblina in Venezuela. It is one of the most variable species in the genus and was once considered to be a variety of H. tatei...

, as well as many infraspecific taxa. Various exploratory expeditions as well as review of existing herbarium specimens has yielded many new species in recent years, mainly through the work of a group of German horticulturalists and botanists (Thomas Carow, Peter Harbarth, Joachim Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

 and Andreas Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

).

Care in cultivation

Heliamphora are regarded by carnivorous plant enthusiasts and experts as one of the more difficult plants to maintain in cultivation. They require cool (the "highland" species) to warm (the "lowland" species) temperatures with a constant and very high humidity
Humidity
Humidity is a term for the amount of water vapor in the air, and can refer to any one of several measurements of humidity. Formally, humid air is not "moist air" but a mixture of water vapor and other constituents of air, and humidity is defined in terms of the water content of this mixture,...

. The highland species, which originate from high in the cloudy Tepui
Tepui
A tepui , or tepuy, is a table-top mountain or mesa found in the Guiana Highlands of South America, especially in Venezuela. The word tepui means "house of the gods" in the native tongue of the Pemon, the indigenous people who inhabit the Gran Sabana....

 mountaintops, include H. nutans, H.ionasi, and H. tatei. The lowland Heliamphora, such as H. minor and H. heterodoxa have migrated to the warmer grasslands at the foot of the Tepuis.

Shredded, long-fibered or live sphagnum moss is preferred as a soil substrate, often with added horticultural lava rock, perlite
Perlite
Perlite is an amorphous volcanic glass that has a relatively high water content, typically formed by the hydration of obsidian. It occurs naturally and has the unusual property of greatly expanding when heated sufficiently...

, and pumice
Pumice
Pumice is a textural term for a volcanic rock that is a solidified frothy lava typically created when super-heated, highly pressurized rock is violently ejected from a volcano. It can be formed when lava and water are mixed. This unusual formation is due to the simultaneous actions of rapid...

. The substrate must always be kept moist and extremely well drained. Misting Heliamphora with purified water
Purified water
Purified water is water from any source that is physically processed to remove impurities. Distilled water and deionized water have been the most common forms of purified water, but water can also be purified by other processes including reverse osmosis, carbon filtration, microfiltration,...

 is often beneficial to maintain high humidity levels.

Propagation through division
Division (horticulture)
Division, in horticulture and gardening, is a method of asexual plant propagation, where the plant is broken up into two or more parts...

 only has a limited rate of success, as many plants that are divided go into shock and eventually die. Germination
Germination
Germination is the process in which a plant or fungus emerges from a seed or spore, respectively, and begins growth. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. However the growth of a sporeling from a spore, for example the...

 of Heliamphora seed is achieved by scattering it on milled sphagnum moss and keeping in bright light and humid conditions. Seed germination begins after many weeks.

Classification

The genus Heliamphora contains the most species in the Sarraceniaceae
Sarraceniaceae
Sarraceniaceae is a family of pitcher plants , belonging to order Ericales .The family comprises three extant genera: Sarracenia , Darlingtonia , and Heliamphora . The extinct Archaeamphora longicervia may also belong to this family...

 family and is joined by the Cobra Lily
Darlingtonia californica
Darlingtonia californica , also called the California Pitcher plant, Cobra Lily, or Cobra Plant, is a carnivorous plant, the sole member of the genus Darlingtonia in the family Sarraceniaceae. It is native to Northern California and Oregon, growing in bogs and seeps with cold running water...

 and the North American pitcher plants
Sarracenia
Sarracenia is a genus comprising 8 to 11 species of North American pitcher plants. The genus belongs to the family Sarraceniaceae, which also contain the closely allied genera Darlingtonia and Heliamphora....

 in that taxon
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...

.

Twenty-three species of Heliamphora are currently recognised, and these are listed in the table below. Unless otherwise stated, all information and taxonomic determinations are sourced from the 2011 work, Sarraceniaceae of South America, authored by Stewart McPherson
Stewart McPherson (geographer)
Stewart R. McPherson is a British geographer.He studied at the University of Durham in England, the University of Tübingen in Germany and Yale University in the United States....

, Andreas Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

, Andreas Fleischmann, and Joachim Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

. Authorities are presented in the form of a standard author citation
Author citation (botany)
In botanical nomenclature, author citation refers to citing the person who validly published a botanical name, i.e. who first published the name while fulfilling the formal requirements as specified by the International Code of Botanical Nomenclature...

, using abbreviations specified by the IPNI. Years given denote the year of the species's formal publication under the current name, thus excluding the earlier basionym
Basionym
Basionym is a term used in botany, regulated by the International Code of Botanical Nomenclature...

 date of publication if one exists.
Species Authority Year Image Distribution Altitudinal distribution
Heliamphora arenicola
Heliamphora arenicola
Heliamphora arenicola is a species of marsh pitcher plant known only from the western side of the Ilu–Tramen Massif in Venezuela's Gran Sabana, where it grows at elevations of less than 2000 m. It may also occur on Karaurin Tepui....

 
Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

, A.Fleischm., Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

 & S.McPherson
Stewart McPherson (geographer)
Stewart R. McPherson is a British geographer.He studied at the University of Durham in England, the University of Tübingen in Germany and Yale University in the United States....

 
2011 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
below 2000 m
Heliamphora ceracea
Heliamphora ceracea
Heliamphora ceracea is a species of marsh pitcher plant known only from the Brazilian side of Pico da Neblina in the Neblina Massif. It has been collected from an elevation of 1900 m....

 
Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

, Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

, Grantsau, Rivadavia, A.Fleischm. & S.McPherson
Stewart McPherson (geographer)
Stewart R. McPherson is a British geographer.He studied at the University of Durham in England, the University of Tübingen in Germany and Yale University in the United States....

 
2011 Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...

 
highland
Heliamphora chimantensis
Heliamphora chimantensis
Heliamphora chimantensis is a species of Marsh Pitcher Plant endemic to Chimanta Tepui in Venezuela. Interestingly, it is more closely related to the southern growing H. tatei and H. neblinae than any of the other species found in the Gran Sabana. All other species known from this region have...

 
Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

, Carow & Harbarth 
2002 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1900–2100 m
Heliamphora ciliata
Heliamphora ciliata
Heliamphora ciliata is a species of Marsh Pitcher Plant endemic to Venezuela. It is restricted to a small number of swampy meadows in the uplands of Gran Sabana. In this habitat it is sympatric with a species of Stegolepis . Unusually for the genus, H. ciliata is a submontane plant, growing at an...

 
Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

, Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

 & A.Fleischm. 
2009 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
900 m
Heliamphora collina
Heliamphora collina
Heliamphora collina is a species of marsh pitcher plant known with certainty only from the Los Testigos Massif in Venezuela, although it may also occur on Ptari Tepui to the east. It grows at elevations of 1700–1825 m....

 
Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

, Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

, S.McPherson
Stewart McPherson (geographer)
Stewart R. McPherson is a British geographer.He studied at the University of Durham in England, the University of Tübingen in Germany and Yale University in the United States....

 & A.Fleischm. 
2011 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1700–1825 m
Heliamphora elongata
Heliamphora elongata
Heliamphora elongata is a species of Marsh Pitcher Plant endemic to Ilu tepui and Tramen Tepui in Venezuela.-References:...

 
Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

 
2004 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1800–2600 m
Heliamphora exappendiculata
Heliamphora exappendiculata
Heliamphora exappendiculata is a species of Marsh Pitcher Plant endemic to the Chimanta massif and Aprada Tepui in Venezuela. It was for a long time considered a variety of H. heterodoxa, but has recently been raised to species rank.-References:* McPherson, S. 2007. Pitcher Plants of the Americas...

 
(Maguire & Steyermark) Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

 & Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

 
2006 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1700–2100 m
Heliamphora folliculata
Heliamphora folliculata
Heliamphora folliculata is a species of Marsh Pitcher Plant endemic to the Aparaman group of tepuis in Venezuela...

 
Wistuba, Harbarth & Carow  2001 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1700–2400 m
Heliamphora glabra
Heliamphora glabra
Heliamphora glabra is a species of Marsh Pitcher Plant native to Serra do Sol in Venezuela. It was for a long time considered a form of H. heterodoxa, but has recently been raised to species rank.-References:...

 
(Maguire) Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

, Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

 & Hoogenstrijd 
2006 borderlands of Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...

, Guyana
Guyana
Guyana , officially the Co-operative Republic of Guyana, previously the colony of British Guiana, is a sovereign state on the northern coast of South America that is culturally part of the Anglophone Caribbean. Guyana was a former colony of the Dutch and of the British...

, and Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1200–2750 m
Heliamphora heterodoxa
Heliamphora heterodoxa
Heliamphora heterodoxa is a species of Marsh Pitcher Plant native to Gran Sabana and the plateau of Ptari Tepui in Venezuela. It was first discovered in 1951 upon Ptari Tepui. This Heliamphora is closely related to H. glabra and the latter was for a long time considered a form of H...

 
Steyerm.
Julian Alfred Steyermark
Julian Alfred Steyermark was an American botanist. His focus was on New World vegetation, and he specialized in the family Rubiaceae.- Life and work :...

 
1951 Guyana
Guyana
Guyana , officially the Co-operative Republic of Guyana, previously the colony of British Guiana, is a sovereign state on the northern coast of South America that is culturally part of the Anglophone Caribbean. Guyana was a former colony of the Dutch and of the British...

?, Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1200–2200 m
Heliamphora hispida
Heliamphora hispida
Heliamphora hispida is a species of Marsh Pitcher Plant endemic to Cerro Neblina, the southernmost tepui of the Guiana Highlands at the Brazil-Venezuela border.-References:...

 
Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

 & Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

 
2000 border between Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...

 and Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1800–3014 m
Heliamphora huberi
Heliamphora huberi
Heliamphora huberi is a species of Marsh Pitcher Plant endemic to the Chimantá massif in Venezuela, where it grows at elevations of 1850–2200 m in a variety of habitats. It has thusfar been recorded from Angasima-tepui, Apacará-tepui, Amurí-tepui, Acopán-tepui, and the border of Torono- and...

 
A.Fleischm., Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

 & Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

 
2009 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1850–2200 m
Heliamphora ionasi  Maguire  1978 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1800–2600 m
Heliamphora macdonaldae
Heliamphora macdonaldae
Heliamphora macdonaldae is a species of Marsh Pitcher Plant endemic to Venezuela. It is closely related to H. tatei and was for a long time considered a form or variety of that species.-References:...

 
Gleason
Henry Gleason
Henry Allan Gleason was a noted American ecologist, botanist, and taxonomist, most recognized for his endorsement of the individualistic/open community concept of ecological succession.- Life and work :...

 
1931 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1500–2300 m
Heliamphora minor
Heliamphora minor
Heliamphora minor is a species of Marsh Pitcher Plant endemic to the Auyan and Chimanta group of tepuis in Venezuela. As the name suggests, it is the smallest species in the genus. It is closely related to H. pulchella....

 
Gleason
Henry Gleason
Henry Allan Gleason was a noted American ecologist, botanist, and taxonomist, most recognized for his endorsement of the individualistic/open community concept of ecological succession.- Life and work :...

 
1939 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1650–2500 m
Heliamphora neblinae
Heliamphora neblinae
Heliamphora neblinae is a species of Marsh Pitcher Plant endemic to Cerro Neblina in Venezuela. It is one of the most variable species in the genus and was once considered to be a variety of H. tatei...

 
Maguire  1978 border between Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...

 and Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
860–2200 m
Heliamphora nutans
Heliamphora nutans
Heliamphora nutans is a species of Marsh Pitcher Plant native to Venezuela, Brazil and Guyana, where it grows on several Tepuis, including Roraima, Kukenan, Tramen and Ilu. H...

 
Benth.
George Bentham
George Bentham CMG FRS was an English botanist, characterized by Duane Isely as "the premier systematic botanist of the nineteenth century".- Formative years :...

 
1840 borderlands of Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...

, Guyana
Guyana
Guyana , officially the Co-operative Republic of Guyana, previously the colony of British Guiana, is a sovereign state on the northern coast of South America that is culturally part of the Anglophone Caribbean. Guyana was a former colony of the Dutch and of the British...

, and Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
2000–2700 m
Heliamphora parva
Heliamphora parva
Heliamphora parva is a species of marsh pitcher plant known only from the Neblina Massif in Venezuela, where it grows at elevations of 1750–2200 m....

 
(Maguire) S.McPherson
Stewart McPherson (geographer)
Stewart R. McPherson is a British geographer.He studied at the University of Durham in England, the University of Tübingen in Germany and Yale University in the United States....

, A.Fleischm., Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

 & Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

 
2011 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1750–2200 m
Heliamphora pulchella
Heliamphora pulchella
Heliamphora pulchella is a species of Marsh Pitcher Plant endemic to the Chimanta group of tepuis in Venezuela. It is one of the smallest species and closely related to H...

 
Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

, Carow, Harbarth & Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

 
2005 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1850–2550 m
Heliamphora purpurascens
Heliamphora purpurascens
Heliamphora purpurascens is a species of marsh pitcher plant known only from the summit area of Ptari Tepui in Venezuela, where it grows at elevations of 2400–2500 m....

 
Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

, A.Fleischm., Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

 & S.McPherson
Stewart McPherson (geographer)
Stewart R. McPherson is a British geographer.He studied at the University of Durham in England, the University of Tübingen in Germany and Yale University in the United States....

 
2011 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
2400–2500 m
Heliamphora sarracenioides
Heliamphora sarracenioides
Heliamphora sarracenioides is a species of Marsh Pitcher Plant native to southern Venezuela, Gran Sabana and Estado Bolivar.-References:...

 
Carow, Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

 & Harbarth 
2005 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
2400–2450 m
Heliamphora tatei
Heliamphora tatei
Heliamphora tatei is a species of Marsh Pitcher Plant endemic to Duida Tepui, Cerro Huachamacari and Marahuaca Tepui in Venezuela. It is closely related to H. neblinae and the latter was at one point considered a variety of H. tatei.-Infraspecific taxa:*Heliamphora tatei var. macdonaldae Maguire ...

 
Gleason
Henry Gleason
Henry Allan Gleason was a noted American ecologist, botanist, and taxonomist, most recognized for his endorsement of the individualistic/open community concept of ecological succession.- Life and work :...

 
1931 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1700–2400 m
Heliamphora uncinata
Heliamphora uncinata
Heliamphora uncinata is a species of Marsh Pitcher Plant endemic to Venezuela. It is known only from the type collection, which was made in a narrow canyon on Amurí-tepui. There it grows at an elevation of approximately 1850 m on sandstone cliff faces in shady conditions. The only other...

 
Nerz
Joachim Nerz
Dr. Joachim Nerz is a taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.-Publications:...

, Wistuba
Andreas Wistuba
Dr. Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.-Publications:...

 & A.Fleischm. 
2009 Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1850 m

Incompletely diagnosed taxa

A further two incompletely diagnosed taxa
Undescribed taxon
In taxonomy, an undescribed taxon is a taxon that has been discovered, but not yet formally described and named. The various Nomenclature Codes specify the requirements for a new taxon to be validly described and named. Until such a description has been published, the taxon has no formal or...

 are known that may represent distinct species in their own right.
Species Distribution Altitudinal distribution
Heliamphora sp. 'Akopán Tepui'
Heliamphora sp. 'Akopán Tepui'
Heliamphora sp. 'Akopán Tepui is an undescribed taxon of marsh pitcher plant known only from Akopán Tepui in Venezuela, where it grows at elevations of 1800–1900 m. It resembles H. heterodoxa in many respects, but has larger and broader pitchers, narrower tepals, and a smaller nectar spoon....

 
Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
1800–1900 m
Heliamphora sp. 'Angasima Tepui'
Heliamphora sp. 'Angasima Tepui'
Heliamphora sp. 'Angasima Tepui is an undescribed taxon of marsh pitcher plant known only from the summit of Angasima Tepui in Venezuela, where it grows at elevations of 2200–2250 m. It resembles H. heterodoxa in many respects, but has a smaller nectar spoon, numerous nectar glands on the...

 
Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 
2200–2250 m

Natural hybrids

At least eleven natural hybrids have been recorded:
  • H. arenicola
    Heliamphora arenicola
    Heliamphora arenicola is a species of marsh pitcher plant known only from the western side of the Ilu–Tramen Massif in Venezuela's Gran Sabana, where it grows at elevations of less than 2000 m. It may also occur on Karaurin Tepui....

     × H. ionasi
  • H. ceracea
    Heliamphora ceracea
    Heliamphora ceracea is a species of marsh pitcher plant known only from the Brazilian side of Pico da Neblina in the Neblina Massif. It has been collected from an elevation of 1900 m....

     × H. hispida
    Heliamphora hispida
    Heliamphora hispida is a species of Marsh Pitcher Plant endemic to Cerro Neblina, the southernmost tepui of the Guiana Highlands at the Brazil-Venezuela border.-References:...

  • H. chimantensis
    Heliamphora chimantensis
    Heliamphora chimantensis is a species of Marsh Pitcher Plant endemic to Chimanta Tepui in Venezuela. Interestingly, it is more closely related to the southern growing H. tatei and H. neblinae than any of the other species found in the Gran Sabana. All other species known from this region have...

     × H. pulchella
    Heliamphora pulchella
    Heliamphora pulchella is a species of Marsh Pitcher Plant endemic to the Chimanta group of tepuis in Venezuela. It is one of the smallest species and closely related to H...

  • H. elongata
    Heliamphora elongata
    Heliamphora elongata is a species of Marsh Pitcher Plant endemic to Ilu tepui and Tramen Tepui in Venezuela.-References:...

     × H. ionasi
  • H. exappendiculata
    Heliamphora exappendiculata
    Heliamphora exappendiculata is a species of Marsh Pitcher Plant endemic to the Chimanta massif and Aprada Tepui in Venezuela. It was for a long time considered a variety of H. heterodoxa, but has recently been raised to species rank.-References:* McPherson, S. 2007. Pitcher Plants of the Americas...

     × H. huberi
    Heliamphora huberi
    Heliamphora huberi is a species of Marsh Pitcher Plant endemic to the Chimantá massif in Venezuela, where it grows at elevations of 1850–2200 m in a variety of habitats. It has thusfar been recorded from Angasima-tepui, Apacará-tepui, Amurí-tepui, Acopán-tepui, and the border of Torono- and...

  • H. exappendiculata
    Heliamphora exappendiculata
    Heliamphora exappendiculata is a species of Marsh Pitcher Plant endemic to the Chimanta massif and Aprada Tepui in Venezuela. It was for a long time considered a variety of H. heterodoxa, but has recently been raised to species rank.-References:* McPherson, S. 2007. Pitcher Plants of the Americas...

     × H. pulchella
    Heliamphora pulchella
    Heliamphora pulchella is a species of Marsh Pitcher Plant endemic to the Chimanta group of tepuis in Venezuela. It is one of the smallest species and closely related to H...

  • H. glabra
    Heliamphora glabra
    Heliamphora glabra is a species of Marsh Pitcher Plant native to Serra do Sol in Venezuela. It was for a long time considered a form of H. heterodoxa, but has recently been raised to species rank.-References:...

     × H. nutans
    Heliamphora nutans
    Heliamphora nutans is a species of Marsh Pitcher Plant native to Venezuela, Brazil and Guyana, where it grows on several Tepuis, including Roraima, Kukenan, Tramen and Ilu. H...

  • H. huberi
    Heliamphora huberi
    Heliamphora huberi is a species of Marsh Pitcher Plant endemic to the Chimantá massif in Venezuela, where it grows at elevations of 1850–2200 m in a variety of habitats. It has thusfar been recorded from Angasima-tepui, Apacará-tepui, Amurí-tepui, Acopán-tepui, and the border of Torono- and...

     × H. pulchella
    Heliamphora pulchella
    Heliamphora pulchella is a species of Marsh Pitcher Plant endemic to the Chimanta group of tepuis in Venezuela. It is one of the smallest species and closely related to H...

  • H. neblinae
    Heliamphora neblinae
    Heliamphora neblinae is a species of Marsh Pitcher Plant endemic to Cerro Neblina in Venezuela. It is one of the most variable species in the genus and was once considered to be a variety of H. tatei...

     × H. parva
    Heliamphora parva
    Heliamphora parva is a species of marsh pitcher plant known only from the Neblina Massif in Venezuela, where it grows at elevations of 1750–2200 m....

  • H. purpurascens
    Heliamphora purpurascens
    Heliamphora purpurascens is a species of marsh pitcher plant known only from the summit area of Ptari Tepui in Venezuela, where it grows at elevations of 2400–2500 m....

     × H. sarracenioides
    Heliamphora sarracenioides
    Heliamphora sarracenioides is a species of Marsh Pitcher Plant native to southern Venezuela, Gran Sabana and Estado Bolivar.-References:...

  • H. sp. 'Akopán Tepui'
    Heliamphora sp. 'Akopán Tepui'
    Heliamphora sp. 'Akopán Tepui is an undescribed taxon of marsh pitcher plant known only from Akopán Tepui in Venezuela, where it grows at elevations of 1800–1900 m. It resembles H. heterodoxa in many respects, but has larger and broader pitchers, narrower tepals, and a smaller nectar spoon....

     × H. pulchella
    Heliamphora pulchella
    Heliamphora pulchella is a species of Marsh Pitcher Plant endemic to the Chimanta group of tepuis in Venezuela. It is one of the smallest species and closely related to H...



Additionally, putative complex hybrids occur on the Neblina Massif
Neblina massif
The Neblina massif is a sandstone mountain located in the Northern Amazon Basin. The massif is a tilted, heavily eroded plateau, with a deep canyon in its central portion , drained by the Baria River...

 among populations of H. ceracea
Heliamphora ceracea
Heliamphora ceracea is a species of marsh pitcher plant known only from the Brazilian side of Pico da Neblina in the Neblina Massif. It has been collected from an elevation of 1900 m....

, H. hispida
Heliamphora hispida
Heliamphora hispida is a species of Marsh Pitcher Plant endemic to Cerro Neblina, the southernmost tepui of the Guiana Highlands at the Brazil-Venezuela border.-References:...

, H. neblinae
Heliamphora neblinae
Heliamphora neblinae is a species of Marsh Pitcher Plant endemic to Cerro Neblina in Venezuela. It is one of the most variable species in the genus and was once considered to be a variety of H. tatei...

, and H. parva
Heliamphora parva
Heliamphora parva is a species of marsh pitcher plant known only from the Neblina Massif in Venezuela, where it grows at elevations of 1750–2200 m....

.
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