The genus ''Heliamphora'' ( or ;

Greek Greek may refer to: 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 of all kno ...

: ''helos'' "

marsh In ecology, a marsh is a wetland that is dominated by herbaceous plants rather than by woody plants.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p More in genera ...

" and ''amphoreus'' "

amphora An amphora (; ; English ) is a type of container with a pointed bottom and characteristic shape and size which fit tightly (and therefore safely) against each other in storage rooms and packages, tied together with rope and delivered by land ...

") contains 24 species of

pitcher plant Pitcher plants are carnivorous plants known as pitfall traps—a prey-trapping mechanism featuring a deep cavity filled with digestive liquid. The traps of pitcher plant are considered to be "true" pitcher plants and are formed by specialized ...

endemic Endemism is the state of a species being found only 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 foun ...

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

.McPherson, S., A. Wistuba, A. Fleischmann & J. Nerz 2011. '' Sarraceniaceae of South America''. 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". The name instead derives from the Greek ''helos'', meaning "marsh", so a more accurate translation of their

scientific name In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...

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, typically insects and other arthropods, and occasionally small mammals and birds. They have adapted to grow in waterlo ...

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. Protein degradation is a major regulatory mechanism of gene expression and contributes substantially to shaping mammalian proteomes. Uncatalysed, the hydrolysis o ...

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

symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...

bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...

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 ...

perennial plant In horticulture, the term perennial (''wikt:per-#Prefix, per-'' + ''wikt:-ennial#Suffix, -ennial'', "through the year") is used to differentiate a plant from shorter-lived annual plant, annuals and biennial plant, biennials. It has thus been d ...

s that grow from a subterranean

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

. ''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 drainage hole (W, E2a, E2b, and E3 Clades) or slit (E1 Clade) in its side that allows excess rainwater to drain away, similar to the overflow on a sink. It was inferred that the drainage structure first evolved as drainage hole in ancestral ''Heliamphora'' populations and further modified into drainage slit in the ancestors of E1 clade. 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. The morphological diversification of ''Heliamphora'' pitchers is both convergent and divergent, likely as a result of adaptive radiation in the geographically complex Guiana highland. 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 catalysis, catalyzes proteolysis, breaking down proteins into smaller polypeptides or single amino acids, and spurring the formation of new protein products ...

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 th ...

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

symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...

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'', ''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 Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...

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 a chemical element; it has Symbol (chemistry), symbol N and atomic number 7. Nitrogen is a Nonmetal (chemistry), nonmetal and the lightest member of pnictogen, group 15 of the periodic table, often called the Pnictogen, pnictogens. ...

and

phosphorus Phosphorus is a chemical element; it has Chemical symbol, symbol P and atomic number 15. All elemental forms of phosphorus are highly Reactivity (chemistry), reactive and are therefore never found in nature. They can nevertheless be prepared ar ...

) 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 member of a family of table-top mountains or mesas found in northern South America, especially in Venezuela, western Guyana, and northern Brazil. The word tepui means "house of the gods" in the native tongue of the ...

s of the Guiana Highlands and their surrounding uplands. Most are found in

Venezuela Venezuela, officially the Bolivarian Republic of Venezuela, is a country on the northern coast of South America, consisting of a continental landmass and many Federal Dependencies of Venezuela, islands and islets in the Caribbean Sea. It com ...

, with a few extending into western

Guyana Guyana, officially the Co-operative Republic of Guyana, is a country on the northern coast of South America, part of the historic British West Indies. entry "Guyana" Georgetown, Guyana, Georgetown is the capital of Guyana and is also the co ...

and northern

Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...

. 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 studie ...

named in 1840 based on a specimen collected by Robert Hermann Schomburgk. 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. Loui ...

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'', 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 and Andreas Wistuba).

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 (meteorology), precipitation, dew, or fog t ...

.Rice, Barry A. (2006). ''Growing Carnivorous Plants''. Timber Press: Portland, Oregon. An amateur botanist in New York City has shown that cultivation of the genus can be achieved with an inexpensive setup consisting of a large plastic crate, a fan, egg cartons, and water bottles filled with ice. 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, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store water, since ...

is preferred as a soil substrate, often with added horticultural lava rock,

perlite Perlite is an amorphous volcanic glass that has a relatively high water content, typically formed by the Hydrate, hydration of obsidian. It occurs naturally and has the unusual property of greatly expanding when heated sufficiently. It is an indu ...

, and

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

. 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 pu ...

is often beneficial to maintain high humidity levels. Propagation through division 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, ...

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 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''; : 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 name and ...

Species

Twenty-four species of ''Heliamphora'' are currently recognized. Unless otherwise stated, all information and taxonomic determinations in the table below are sourced from the 2011 work '' Sarraceniaceae of South America'' authored by Stewart McPherson, Andreas Wistuba, Andreas Fleischmann, and Joachim Nerz. 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 in ...

. 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 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 hybrids 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 can ...

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.

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 *
''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 Taxa named by George Bentham