Pinguicula

Pinguicula ramosa

The fat herbs ( Pinguicula ), called in Liechtenstein and parts of the German-speaking Switzerland and fat leaves, are a genus of carnivorous plants ( carnivores ) from the family of water hose plants ( Lentibulariaceae ). Currently, about 85 species are known, of which four are endemic in German-speaking, the majority found, however, in Central America.

  • 6.1 subgenus Isoloba 6.1.1 Section Agnata
  • 6.1.2 Section Cardiophyllum
  • 6.1.3 Section Disco Radix
  • 6.1.4 Section heterophyllum
  • 6.1.5 Section Isoloba
  • 6.2.1 Section Crassifolia
  • 6.2.2 Section Homophyllum
  • 6.2.3 Section Longitubus
  • 6.2.4 Section Nana
  • 6.2.5 Section Orcheosanthus
  • 6.2.6 Section Orchidioides
  • 6.2.7 Section Pinguicula
  • 6.3.1 Section Ampullipalatum
  • 6.3.2 Section micranthus
  • 6.3.3 Section Temnoceras

Features

Almost all fat herbs are perennial herbaceous plants (except for the annual species P. crenatiloba, P. sharpii, P. and P. pumila takakii ). All species form indigenous rosettes.

Growth forms

Coarse, the fatty herbs are divided into two main groups, each with two sub-groups. The subdivision of the main groups goes back on climatic conditions, the. Subgroups morphological features

Generally for all Butterworts that they form so-called seasonal alternating vegetative and generative rosettes. Tropical growth habits stop doing any rosette formation with the formation of a flower and go to this again in the next rosette shape over. Although the so-called " tempered growth forms ", however, form a vegetative rosette, but this does not bloom, but forms at its center a Hibernakel, which serves as a wintering organ. After the formation of Hibernakels the leaves of the vegetative rosette die. Only after overwintering and new leaves of a rosette, the plant begins the formation of a flower.

The distinguishing feature of the sub-groups, in turn, serves the shape of rosettes of a kind When vegetative and generative rose a sort are identical in appearance, it is called " homophyllem growth " if they differ in shape and / or size, this is called " heterophyllem growth ".

Thus results in four groups of forms:

  • Tropical habit type: Species that do not know any cold periods and variable form rosettes and blossom. Tropical - heterophyller growth habit Type: vegetative and generative rose a kind differ in shape and / or size. Examples: P. cyclosecta, P. acuminata or P. moranensis.
  • Tropical - homophyller growth habit Type: vegetative and generative rose a sort are identical in appearance. Examples: P. emarginata, P. filifolia or P. pumila.
  • Temperate growth habit Type: species in temperate zones with cold spells. They form alternating rosettes and blossom and winter changing. Temperate - heterophyller growth habit Type: vegetative and generative rose a kind differ in shape and / or size. Examples: P. lutea or P. lusitanica.
  • Temperate - homophyller growth habit Type: vegetative and generative rose a sort are identical in appearance. Examples: P. alpina, P. grandiflora or P. vulgaris.

Root

The root system of Butterworts is weak. It is white and fine and serves primarily anchored in the ground to absorb moisture and trace elements. In species of temperate zones, which form Hibernakel, they are rootless, with the exception of the Alpine - fat herb. In the (few ) epiphytic species (eg P. lignicola ) the roots are additionally provided with adhesive discs.

The leaves are smooth, firm and succulent, bright green of mostly, sometimes reddish tinged color. They are - depending on the type - 2-30 inches tall. In relation to the overall size of the plant, the leaves are large and each shaped differently by clan; However, often found ovate, spatelige or elongated shapes.

Like all species of the family of water hose plants are carnivorous fat herbs. For secretion of the leaf surface with two types of glands is provided: stalked glands and sessile glands. The secreted by the stalked glandular secretion catch, stick to the prey, makes the leaves shine greasy. This gloss to draw more - similar to other carnivores with sticky traps ( sundew, rainbow plants) - the prey. The captured animal device then in contact with the seated glands of the surface of which subsequently the enzyme-containing digestive juice is separated. Proven enzymes are amylase, esterase, phosphatase, protease and ribonuclease. The leaves are movable, albeit very slowly, and can be rolled up at the edges about not exploiting dividend in the center of the sheet prey. They can also form small depressions in the leaf surface to the prey, that collects digestive juices for better decomposition of the prey. Journal of areas that have already been made ​​loot, it can not decompose once more prey, as the stalked glands are only " one-way" glands. The dissolved by the decomposition of the prey nutrients are then absorbed through openings in the cuticle. Since the cuticle protects the plants from drying out and this is broken in fat herbs through the openings in many cases, the species is dependent on humid habitats.

The prey spectrum usually comprises smallest flying insects, but the plants can also efficiently utilize pollen that is blown on their relatively large leaf surface.

Flowers

As with almost all carnivores, the hermaphrodite flowers protrude far beyond the actual plant to an endeavor to prevent from potential pollinators. The solitary and long blooming flowers, in which the relationship of fat herbs with the other water hose plants shows most clearly, are zygomorphic, the crown has two pronounced lips and spurred on their approach. The deformed calyx is usually five-piece, the upper lip three -, two-piece lower lip. Usually the flowers are blue, violet or white, often with yellow, greenish or reddish stains are rare. The latter is particularly pronounced in the strongly red flowers of P. laueana and P. caryophyllacea.

The exact shape of the flower is used as a diagnostic feature for the subdivision of the genus into subgenera, as well as the flower color, with multicolored flowers and their combination.

Fruit and seeds

The egg-shaped to spherical capsule fruits open with drought in two flaps and close when it absorbs moisture again in order to protect and promote their dispersal by wind scattering as the numerous, fine, between 0.5 and 1 millimeter long, golden-brown seeds. The seeds of most species bear on their surface a grid pattern in order to thus drive on water surfaces without sinking can (except epiphythische species whose seed surface is smooth ). The chromosome number of fat herb has two bases, either x = 8 or x = 11, an exception to this rule is P. lusitanica, the basic number is x = 6.

Vegetative propagation

In addition to the generative seed propagation, there are several ways of vegetative reproduction in this genus. Many species of the genus ( eg P. vulgaris ) form at the time of flowering or shortly afterwards small bulbils from which to develop new, genetically identical plants. A few species also form processes ( eg P. calyptrata, P. vallisnerifolia ) or Kindeln ( eg P. heterophylla, P. primuliflora ) at their tips.

Dissemination

Butterworts are distributed over the entire northern hemisphere of the earth; in Africa, the spread is limited to the far northwest. Above all, the plants are, however, found in Central and South America, where the occurrence radiate far across the equator to Tierra del Fuego. Not represented is the only genus in Australia and New Zealand.

The area of ​​origin of Pinguicula is probably the present-day Central America, which also forms the center of diversity of the genus - around 50 percent of the approximately eighty species are found in Mexico alone, there are twelve species in Europe, in North America ten. From Mexico, the forerunner of today's fat herbs have probably spread to the Europe of today, before the former supercontinent drifted apart unbridgeable (see also: Laurasia, Pangaea ).

Far more than half of all species has very limited distribution areas. The two most common types, namely the Common fat (Pinguicula vulgaris ) and the Alps - fat (Pinguicula alpina), are native to the German-speaking countries. As an alpine plant is also found in Austria and Switzerland nor the Dünnspornige fat (Pinguicula leptoceras ) and in Switzerland, the Large-flowered fat (Pinguicula grandiflora ), for the latter is also a angesalbter location in North Rhine -Westphalia known.

Habitat requirements

In general Butterworts prefer nutrient-poor, neutral to alkaline soil as locations. Different types but other soils such as acidic peat have conquered ( P. vulgaris, P. calyptrata, P. lusitanica ) or pure gypsum ( P. gypsicola ) or partly settle on bare, upright rock ( P. ramosa, P. vallisneriifolia ). A few species have also changed to epiphytes ( P. casabitoana, P. lignicola ).

Butterworts need biotopes, the wet at least for the growing season to be wet, but can in the form of succulent winter rosettes times relative to extremely severe drought survive. As moisture may well be sufficient spray or leachate. Unlike many other carnivores Butterworts prefer indeed open, but not directly sunlit locations, often even semi -shady to shady locations are preferred.

Endangering

The threat of Pinguicula can not bring to a common denominator, as the situation of the species is very different. They differ greatly in their distribution and have differing habitat requirements. However, particularly at high risk are some endemic species such as P. ramosa, P. casabitoana or P. fiorii, where as most threatening factor in this habitat destruction is mentioned. The North American P. ionantha is listed in Appendix 1 of CITES. The species occurring in Germany are all protected by Federal Species Protection Ordinance.

Botanical history

The fat herb was first mentioned as " zitroch chrawt or smalz chrawt " by Vitus Auslasser in his herbal book " Macer de Herbarium " from 1479. The name " Zittrochkraut " is also still in use today for the Butterworts in Tyrol. The botanical name comes from Conrad Gesner back, who in his " Horti Germaniae " referred in 1561 to the fat glossy leaves, " propter pinguia et tenera folia ... " (Latin pinguis = bold). The current name " fat herb" or rarely " Bacopa " reflects this origin name.

1583 differed Clusius in his " Historia stirpium rariorum by Pannoniam, Austriam " already two forms, a blue flowering ( = P. vulgaris ) and a white-flowering ( = P. alpina); 1753 took Linnaeus in his " Species Plantarum " in addition P. villosa and P. lusitanica on. Especially by new discoveries in the exploration of new continents, the number of known species increased in the 19th century quickly; 1844 were already 32 species known.

As W. Marshall Charles Darwin made ​​in 1875 on the on the leaves stick insects lasting attention, these plants examined more closely and found the carnivory of the genus. Siegfried Jost Casper difference in his pioneering monograph of the genus from 1966 still only 46 species, a number that nearly doubled in the next four decades. Other species are mainly in Mexico still found, but also in Europe, some species have been first described only in the recent past, so now are over 80 known species.

An important step to further explore the genre was the establishment of the International Pinguicula Study Group ( IPSG ) in the 1990s, an organization of scientists and amateurs alike.

System

Within the family of water hose plants ( Lentibulariaceae ) apply the herbal fat than the type with the most original features. Systematically, they are divided by Siegfried Jost Casper into three subgenera with 15 sections. Although a phylogenetic study of the genus ( more under phylogenetics ) made ​​the need for a revision of the genus systematics clear Caspers system is, however, still in use.

Subgenus Isoloba

Section Agnata

  • Pinguicula Agnata
  • Pinguicula albida
  • Pinguicula filifolia
  • Pinguicula gigantea
  • Pinguicula benedicta
  • Pinguicula cubensis
  • Pinguicula infundibuliformis
  • Pinguicula pilosa
  • Pinguicula lippoldii
  • Pinguicula toldensis

Section Cardiophyllum

  • Pinguicula crystallina

Section Disco Radix

  • Pinguicula casabitoana
  • Pinguicula lignicola

Section heterophyllum

  • Pinguicula acuminata
  • Pinguicula heterophylla
  • Pinguicula kondoi
  • Pinguicula mirandae
  • Pinguicula parvifolia
  • Pinguicula rotundiflora
  • Pinguicula imitatrix
  • Pinguicula conzattii

Section Isoloba

Subgenus Pinguicula

Section Crassifolia

  • Pinguicula esseriana
  • Pinguicula ehlersiae
  • Pinguicula debbertiana
  • Pinguicula jaumavensis

Section Homophyllum

  • Pinguicula greenwoodii
  • Pinguicula jackii

Section Longitubus

  • Pinguicula calderoniae
  • Pinguicula crassifolia
  • Pinguicula hemiepiphytica
  • Pinguicula laueana
  • Pinguicula utricularioides

Section Nana

  • Pinguicula villosa

Section Orcheosanthus

  • Pinguicula macrophylla
  • Pinguicula oblongiloba
  • Pinguicula stolonifera
  • Pinguicula colimensis
  • Pinguicula cyclosecta
  • Pinguicula mesophytica
  • Tailed fat (Pinguicula moranensis )
  • Pinguicula orchidioides
  • Pinguicula potosiensis
  • Pinguicula rectifolia
  • Pinguicula zecheri
  • Pinguicula gypsicola
  • Pinguicula moctezumae
  • Pinguicula elizabethiae

Section Orchidioides

Section Pinguicula

  • Pinguicula balcanica
  • Pinguicula corsica
  • Pinguicula poldinii
  • Großblütiges fat (Pinguicula grandiflora )
  • Dünnsporniges fat (Pinguicula leptoceras )
  • Pinguicula longifolia
  • Pinguicula macroceras
  • Pinguicula mundi
  • Pinguicula nevadensis
  • Pinguicula vallisneriifolia
  • Mean fat (Pinguicula vulgaris)
  • Pinguicula mariae
  • Pinguicula Apuana

Subgenus Temnoceras

Section Ampullipalatum

  • Pinguicula antarctica
  • Pinguicula calyptrata
  • Pinguicula chilensis
  • Pinguicula chuquisacensis
  • Pinguicula involuta
  • Pinguicula elongata

Section micranthus

  • Pinguicula algida
  • Alps - fat (Pinguicula alpina)
  • Pinguicula ramosa
  • Pinguicula variegata

Section Temnoceras

  • Pinguicula clivorum
  • Pinguicula crenatiloba
  • Pinguicula emarginata
  • Pinguicula gracilis
  • Pinguicula immaculate

Phylogenetics

Phylogenetic analysis proved that all subgenera and many of the species -rich sections ( with a total of more than 60 percent of affected species ) are polyphyletic or paraphyletic. To keep the cladogram clearly, we will refrain from a too detailed reproduction; polyphyletic sections are marked in the sequence with *.

Clade I ( sections Temnoceras *, * Orcheosanthus, Longitubus, heterophyllum *, Agnata *, * Isoloba, Crassifolia )

Clade II ( section micranthus * = P. alpina)

Clade III ( sections micranthus *, Nana )

Clade IV ( section Pinguicula )

Clade V ( sections Isoloba *, Ampullipalatum, Cardiophyllum )

Use

Fat herbs are among lovers of carnivores widespread ornamental plants; especially species of temperate zones and easier culturable Mexican species are often found in corresponding collections. The most widespread are the two hybrids Pinguicula × ' Sethos ' and Pinguicula × 'Weser'. Both are crosses of Pinguicula Pinguicula ehlersiae with moranensis, which are also used by commercial orchid growers for pest control.

Asterids Middle America (region) Conrad Gessner Species description Integrated Authority File
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