The genus ''Heliamphora'' ( or ;

Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...

: ''helos'' "

marsh A marsh is a wetland that is dominated by herbaceous rather than woody plant species.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p Marshes can often be found at ...

" and ''amphoreus'' "

amphora An amphora (; grc, ἀμφορεύς, ''amphoreús''; English plural: amphorae or amphoras) is a type of container with a pointed bottom and characteristic shape and size which fit tightly (and therefore safely) against each other in storag ...

") contains 23 species of

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 ...

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

.McPherson, S., A. Wistuba, A. Fleischmann & J. Nerz 2011. ''

Sarraceniaceae of South America ''Sarraceniaceae of South America'' is a monograph on the pitcher plants of the genus ''Heliamphora'' by Stewart McPherson, Andreas Wistuba, Andreas Fleischmann, and Joachim Nerz. It was published in September 2011 by Redfern Natural History Prod ...

''. Redfern Natural History Productions, Poole. 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 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 ...

would be marsh pitcher plants. Species in the genus ''Heliamphora'' are

carnivorous plant Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryot ...

s that consist of a modified leaf form that is fused into a tubular shape. They have evolved mechanisms to attract, trap, and kill insects; and control the amount of water in the pitcher. At least one species ('' H. tatei'') produces its own

proteolytic Proteolysis is the breakdown of proteins into smaller polypeptides or amino acids. Uncatalysed, the hydrolysis of peptide bonds is extremely slow, taking hundreds of years. Proteolysis is typically catalysed by cellular enzymes called proteases, ...

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

symbiotic Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...

bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...

Morphology

All ''Heliamphora'' species are

herbaceous Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition of t ...

perennial plant A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...

s that grow from a subterranean

rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...

. ''Heliamphora'' species form stemless rosettes and leaf height ranges from a few centimeters ('' H. minor'', '' H. pulchella'') up to more than ('' H. ionasi'', '' H. tatei''). ''Heliamphora'' possess tubular traps formed by rolled leaves with fused edges. Marsh pitcher plants are unusual among

s in that they lack lids ( opercula), 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. ASRHeli

Carnivory

Though often counted among the various

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 A protease (also called a peptidase, proteinase, or proteolytic enzyme) is an enzyme that catalyzes (increases reaction rate or "speeds up") proteolysis, breaking down proteins into smaller polypeptides or single amino acids, and spurring the ...

ribonuclease Ribonuclease (commonly abbreviated RNase) 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 ...

phosphatase In biochemistry, a phosphatase is an enzyme that uses water to cleave a phosphoric acid Ester, monoester into a phosphate ion and an Alcohol (chemistry), alcohol. Because a phosphatase enzyme catalysis, catalyzes the hydrolysis of its Substrate ...

s, ''etc.''), relying instead on the enzymes of

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 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 ...

. Field studies of '' H. nutans'', ''H. heterodoxa'', ''H. minor'', and ''H. ionasi'' have determined that none of these species produce their own

enzymes.Jaffe, K., Michelangeli, F., Gonzalez, J.M., Miras, B., and Ruiz, M.C. (1992). Carnivory in Pitcher Plants of the Genus ''Heliamphora'' (Sarraceniaceae). ''New Phytologist'', 122(4): 733-744. (First page available online
JSTOR PDF of first page and HTML text of abstract
/ref> ''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 Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...

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

and

phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...

) 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

to the

tepui A tepui , or tepuy (), is a table-top mountain or mesa found in South America, especially in Venezuela and western Guyana. The word tepui means "house of the gods" in the native tongue of the Pemon, the indigenous people who inhabit the Gran S ...

s of the

Guiana Highlands The Guianas, sometimes called by the Spanish loan-word ''Guayanas'' (''Las Guayanas''), is a region in north-eastern South America which includes the following three territories: * French Guiana, an overseas department and region of France * G ...

and their surrounding uplands. Most are found in

Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivariana de Venezuela), is a country on the northern coast of South America, consisting of a continental landmass and many islands and islets in th ...

, with a few extending into western

Guyana Guyana ( or ), officially the Cooperative Republic of Guyana, is a country on the northern mainland of South America. Guyana is an indigenous word which means "Land of Many Waters". The capital city is Georgetown. Guyana is bordered by the ...

and northern

Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...

. 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 (22 September 1800 – 10 September 1884) was an English botanist, described by the weed botanist Duane Isely as "the premier systematic botanist of the nineteenth century". Born into a distinguished family, he initially studi ...

named in 1840 based on a specimen collected by

Robert Hermann Schomburgk Sir Robert Hermann Schomburgk (5 June 1804 – 11 March 1865) 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 missio ...

. This remained the only known species until Henry Allan Gleason described ''H. tatei'' and '' H. tyleri'' in 1931, also adding ''H. minor'' in 1939. Between 1978 and 1984,

Julian Alfred Steyermark Julian Alfred Steyermark (January 27, 1909 – October 15, 1988) was a Venezuelan American botanist. His focus was on New World vegetation, and he specialized in the family Rubiaceae. Life and work Julian Alfred Steyermark was born in St. Louis, ...

and

Bassett Maguire Bassett Maguire (August 4, 1904 – February 6, 1991) was an American botanist, head curator of the New York Botanical Garden, and a leader of scientific expeditions to the Guyana Highlands in Brazil and Venezuela. Life Maguire was born in Gadsd ...

revised the genus (to which Steyermark had added ''H. heterodoxa'' in 1951) and described two more species, ''H. ionasi'' and '' H. neblinae'', as well as many

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 ...

. 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 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. ...

and

Andreas Wistuba Andreas Wistuba (born 4 March 1967) 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. Pub ...

Care in cultivation

''Heliamphora'' are regarded by carnivorous plant enthusiasts and experts as one of the more difficult plants to maintain in cultivation. The genus requires cool (the "highland" species) to warm (the "lowland" species) temperatures with a constant and very high

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

.Rice, Barry A. (2006). ''Growing Carnivorous Plants''. Timber Press: Portland, Oregon. The highland species, which originate from high on the humid

mountaintops, include ''H. nutans'', ''H. ionasi'', and ''H. tatei''. The lowland ''Heliamphora'', such as ''H. ciliata'' and ''H. heterodoxa'' have migrated to the warmer grasslands at the foot of the tepuis. Shredded, long-fibered, or live

sphagnum moss ''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 ...

is preferred as a soil substrate, often with added horticultural

lava rock Volcanic rock (often shortened to volcanics in scientific contexts) is a rock formed from lava erupted from a volcano. In other words, it differs from other igneous rock by being of volcanic origin. Like all rock types, the concept of volcanic r ...

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. It is an industrial m ...

, and

pumice Pumice (), called pumicite in its powdered or dust form, is a volcanic rock that consists of highly vesicular rough-textured volcanic glass, which may or may not contain crystals. It is typically light-colored. Scoria is another vesicular vol ...

. The substrate must always be kept moist and extremely well drained. Misting ''Heliamphora'' with

purified water Purified water is water that has been mechanically filtered or processed to remove impurities and make it suitable for use. Distilled water was, formerly, the most common form of purified water, but, in recent years, water is more frequently puri ...

is often beneficial to maintain high humidity levels. Propagation through

division Division or divider may refer to: Mathematics *Division (mathematics), the inverse of multiplication *Division algorithm, a method for computing the result of mathematical division Military *Division (military), a formation typically consisting ...

only has a limited rate of success, as many plants that are divided go into shock and eventually die.

Germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fer ...

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 are a family of pitcher plants, belonging to order Ericales (previously Nepenthales). The family comprises three extant genera: ''Sarracenia'' (North American pitcher plants), '' Darlingtonia'' (the cobra lily or California pitch ...

family and is joined by the cobra lily (''Darlingtonia californica'') and the North American pitcher plants (''Sarracenia'' spp.) in that

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 ...

Species

Twenty-three species of ''Heliamphora'' are currently recognized. Unless otherwise stated, all information and taxonomic determinations in the table below are sourced from the 2011 work ''

'' authored by Stewart McPherson,

, Andreas Fleischmann, and

. Authorities are presented in the form of a standard author citation, using abbreviations specified by the

IPNI 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 ...

. Years given denote the year of the species's formal publication under the current name, not the earlier

basionym In the scientific name of organisms, basionym or basyonym means the original name on which a new name is based; the author citation of the new name should include the authors of the basionym in parentheses. The term "basionym" is used in both botan ...

date of publication if one exists.

Incompletely diagnosed taxa

A further two incompletely diagnosed taxa are known that may represent distinct species in their own right.

Varieties

Two

varieties Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...

are currently recognised in the genus: ''H. minor'' var. ''pilosa'' and ''H. minor'' var. ''minor''. Additionally, an undescribed variant of '' H. pulchella'' , with traps lacking long retentive hairs is known from Amurí Tepui.

Cultivars

There are currently four registered Heliamphora cultivars including Heliamphora 'Cyclops' (A. Smith), Heliamphora 'Patasola' (B. Tincher & J. Lei), ''Heliamphora'' 'Red Mambo' (F. Boulianne), and Heliamphora 'Scylla' (I. Bogdanow).

Natural hybrids

At least eleven

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: *'' H. arenicola'' × '' H. ionasi'' *'' H. ceracea'' × '' H. hispida'' *'' H. chimantensis'' × '' H. pulchella'' *'' H. elongata'' × '' H. ionasi'' *'' H. exappendiculata'' × '' H. huberi'' *'' H. exappendiculata'' × '' H. pulchella'' *'' H. glabra'' × '' H. nutans'' *'' H. huberi'' × '' H. pulchella'' *'' H. neblinae'' × '' H. parva'' *'' H. purpurascens'' × '' H. sarracenioides'' * ''H.'' sp. 'Akopán Tepui' × '' H. pulchella'' Additionally, putative complex hybrids occur on the

Neblina Massif Cerro de la Neblina (lit. "Mountain of the Mist"), also known as Serra da Neblina in Brazil and Sierra de la Neblina in Venezuela, is a sandstone massif located in the northern Amazon Basin. It is a tilted, heavily eroded plateau, with a deep ca ...

among populations of '' H. ceracea'', '' H. hispida'', '' H. neblinae'', and '' H. parva''. Putative crosses between ''H. macdonaldae'' and ''H. tatei'' have also been recorded in the southern part of

Cerro Duida Cerro Duida, named Yennamadi by the Ye'kuana, ye'kwana, is a very large tepui in Amazonas, Venezuela, Amazonas state, Venezuela. It has an uneven and heavily inclined plateau, rising from highs of around in the north and east to a maximum of on ...

Phylogeny and Diversification

Closely related species tend to be geographically closely distributed. Major ''Heliamphora'' clades probably emerged through both geographical separation and dispersal in the Guiana Highlands during Miocene with more recent diversification driven by vertical displacement during the Pleistocene glacial-interglacial thermal oscillations. Heli phylo

References

External links

Growing
an
propagating ''Heliamphora''
from the

International Carnivorous Plant Society The International Carnivorous Plant Society (ICPS) is a non-profit organization founded in 1972. It is the International Cultivar Registration Authority for carnivorous plants. As of June 2011, the society had around 1400 members. The ICPS is prob ...

*
''Heliamphora'' FAQs
from Barry Rice's Carnivorous Plant FAQ {{Taxonbar, from=Q1758059 Ericales genera Carnivorous plants of South America Guayana Highlands Flora of the Tepuis