Heliamphora

Heliamphora chimantensis ( in culture)

The marsh pitchers ( Heliamphora ) are a präkarnivore genus of the family of Pitcher plants ( Sarraceniaceae ).

• 2.1 Karnivorie

Description

Vegetative habit

The species of the genus Heliamphora are clump-forming, perennial, herbaceous plants growing. Up to growing in the form of up to four meters high perennials Heliamphora tatei make all kinds indigenous rosettes. The simple or branched, robust rhizome horizontal stems and grows upward so incur additional growth points. Starting from a single plant may require extensive clumps of thousands of individuals arise that are several square meters in size so (especially in Heliamphora chimantensis ) for decades. The fibrous brown roots reach a length of up to 25 centimeters.

All species have an age-dependent Blattdimorphismus, which means that the leaves of a young plant significantly from those of an older differ in shape. The young leaves are shaped approximately the same for all species; the species-specific features are formed only after up to 4 years.

The three to ten completely upright to slightly to the center of the rosette curved, funnel-shaped or cylindrical leaves are falling from the pit type and reach a height of about 8 to 10 centimeters in Heliamphora minor and Heliamphora pulchella than 20 cm at Heliamphora nutans up to about half a meters at Heliamphora ionasii.

The traps have on the outer edge of the peristome ( = the boundary of the tube opening ) a very small, often contrasting colored, ear- or helmet-shaped opening on the hose bent upright lid rudiment. An exception is the type Heliamphora sarracenioides whose lid similar to a roof covers the entire opening laterally and perpendicular to the case inside and only a small opening to the middle rosette leaves something. The outside of the traps is hairless, the inside of the funnel is partially covered with hair downward, making it difficult to escape from inside fallen prey. The shape of the traps prevents Fly Out fallen into insects, since a downward spiral caused by the wing motion in the traps. At the front of the tube, the tubes have a so-called Ala, a wing which is densely covered with nectaries. Halfway up the Ala there is a small opening can drain through the excess rainwater, a little hair in it acts as a screen and prevents swept out to become prey.

Flowers, fruits and seeds

On the long, upright flower stems, the few flowers are in a cluster. The mostly four, rarely five or six, acuminate - ovate petals are white ( sometimes tinged with green ) to red, in some species, they turn over the heyday of white to red. The ovary is hairy upper constant and dense. Around him grow ten to twenty stamens, it's just a pen available.

Ecology

Karnivorie

Ways of attracting insects are on the one hand the striking coloration of the leaves, especially the cover rudiments and on the other hand, the reflection of the UV light alone through the inner leaf surface, the case opening clearly stands for insect eyes against the environment and the leaf outside. This is supported by the secretion of nectar by nectar glands at the Ala as well as particularly strong on the cover rudiment, to which further chemical attractants are excreted. The hoses can be divided into four zones divide: Zone 1 is the cover rudiment, Zone 2, the top tube section and the zones 3 and 4, each deeper sections of the tube. Each of these zones has thereby the prey capture a specific function for which it is accordingly equipped differently. The first zone, the densely populated with nectary cover rudiment, separates larger quantities of strongly scented nectar from and Sarracenin ( an antimicrobial and predators abhaltendes iridoid ) Cineron ( an insecticide precursor), more than insect pheromones effective ester, phenyl acetaldehyde ( a chemical for communication among insects), erucic amide ( a lubricant, on the one hand, the secretion holds liquid and at the same time to catch the insects favors ), numerous alkanes and xylene. These substances are often the attraction or the catching of prey insects, but may also have other, such as numbing or adhesive effects.

Insects visiting the plants, aiming to cover rudiment ( soil insects along the Ala, flying insects immediately ) and rush out of there either in the hose or of other nectaries to the hose inside. So they get into the densely populated with downward facing hairs Zone 1, which is just more like an island filled with nectar glands. Since the downward pointing hairs block the way back to the edge, the insect can only move further downwards until it reaches the abrupt beginning of the approach zone 3, whose surface is hairless and smooth, so that the animal slides down the tube and into the liquid falls from zone 4, where the digestion takes place.

Only by a Sumpfkrugart, Heliamphora tatei, the production of enzymes has been able to be detected by the plants themselves, even here, however, only in closed and young tubes. Tests in some other species were unsuccessful. Thus ( with just the exception of Heliamphora tatei ) the species is not classified as a carnivore in the strict sense, but as " präkarnivor ". In what way exactly does the digestion of prey, has not yet been secured, two hypotheses are assumed to be likely: first, could a resident of the liquid bacterial fauna decompose the prey. Secondly, a so-called autolysis, ie a self-dissolution of the booty would be possible, because the majority of the prey of the marsh pitchers is 80 to 95% of ants with an often predatory nature, bring the digestive enzymes in their interior. Because enzymes have a catalytic effect, so do not consume and also do not decompose, they might accumulate and thus serve to digest other prey, which in turn bring again enzymes. It is also possible that several of these factors interact.

In addition to ants ( preferably of the genus Linepithema, Camponotus and Solenopsis ) prey on marsh pitchers significant fractions also beetles, mosquitoes, fungus gnats, Diptera, Hymenoptera, and occasionally larger prey such as butterflies, scorpions or stoneflies.

Marsh pitchers form many of the characteristics of their Karnivorie exclusively to extremely light-intensive locations, the overshadowing by vegetation or landscape features leads not only to a relatively slow growth rate and the absence of nectar, the strong coloration, the hair and only weak cover rudiments. This has led to the assumption that marsh pitchers need the Karnivorie only in situations increased nutrient requirements ( eg in the case of rapid growth through optimal site conditions ) as an additional source of nutrients.

Dissemination

The genus is almost exclusively on the so-called tepuis, which are difficult to access mesas on the plateau on the border between Brazil, Venezuela and Guyana, native. Only Heliamphora Heliamphora sarracenioides heterodoxa and can also be found in the lower elevations of this region.

Evolution

For a long time Heliamphora was kept because of their simple construction case for one of the most primitive carnivorous plants. Recent genetic and biological studies lay flowers but suggests that Heliamphora probably has a long history of development and adopted its simple form only recently.

System

By 2010, 19 species have been described, so are the pitchers swamp the largest genus of Pitcher plants. The exact taxonomy and ecology of this species is not yet clarified beyond doubt; especially the status of Heliamphora Heliamphora tyleri and macdonaldae is controversial, with now penetrated to the status of Heliamphora tyleri the view of synonymy to Heliamphora tatei.

• Heliamphora chimantensis Wistuba, Carow & Harbarth
• Heliamphora ciliata Wistuba, mink & A. Fleischm.
• Heliamphora elongata mink
• Heliamphora exappendiculata ( Maguire & Steyerm. ) Mink & Wistuba
• Heliamphora folliculata Wistuba, Harbarth & Carow
• Heliamphora glabra ( Maguire ) mink, Wistuba & Hoogenstr.
• Heliamphora heterodoxa Steyerm.
• Heliamphora hispida Wistuba & Mink
• Heliamphora huberi A.Fleischm. , Wistuba & Mink
• Heliamphora ionasii Maguire
• Heliamphora macdonaldae Gleason
• Heliamphora minor Gleason
• Heliamphora neblinae Maguire
• Heliamphora nutans Benth.
• Heliamphora pulchella Wistuba, Carow, Harbarth & Mink
• Heliamphora sarracenioides Carow, Wistuba & Harbarth
• Heliamphora tatei Gleason
• Heliamphora tyleri Gleason
• Heliamphora uncinata mink, Wistuba & A.Fleischm.

Botanical history

The type species is Heliamphora nutans, which was first described in 1840 by George Bentham based on a collected by Robert Hermann Schomburgk copy the genus name is derived from the ancient Greek names for " marsh " ( ἕλος effortlessly ) and " jug " ( άμφορεύς amphoreus ) composed. In English-speaking the generic name has sometimes been erroneously translated as "Sun Pitcher " (from the Greek helios ).

Over ninety years, the genus was monotypic before Henry Allan Gleason 1931 Heliamphora tatei, Heliamphora Heliamphora minor described tyleri and 1939. Between 1978 and 1984, edited Julian Alfred Steyermark (who had already in 1951 the genus Heliamphora to heterodoxa supplemented ) and Bassett Maguire, the genus neblinae further described and in addition to numerous infraspecific taxa and two new species, namely Heliamphora ionasii and Heliamphora. Since 1999 - a result of multiple expeditions and the sighting of existing herbarium - especially a group of German researchers, consisting of Thomas Carow, Peter Harbarth, Joachim mink, Andreas Wistuba and Andreas Fleischmann to made ​​worthy of the genus and continuously described new species, the description of other taxa is still pending.

Swell

• Barthlott, Wilhelm; Porembski, Stefan; Seine, Rüdiger; Theisen, Inge: carnivores, Stuttgart, 2004, ISBN 3-8001-4144-2
• Lloyd, Francis Ernest: The Carnivorous Plants, 1942
• McPherson, Stewart: Pitcher Plants of the Americas, Blacksburg, 2007, ISBN 0-9399-2374-2
• Jaffe, K., Michelangeli, F., Gonzalez, JM, Miras, B., and Ruiz, MC: Carnivory in Pitcher Plants of the genus Heliamphora ( Sarraceniaceae ), in: New Phytologist, vol 122, no. 4, 1992, pp. 733-744