# Pinguicula

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Genus of flowering plants in the family Lentibulariaceae

Not to be confused with [Pinguecula](/source/Pinguecula).

Pinguicula Pinguicula moranensis Scientific classification Kingdom: Plantae Clade: Embryophytes Clade: Tracheophytes Clade: Spermatophytes Clade: Angiosperms Clade: Eudicots Clade: Asterids Order: Lamiales Family: Lentibulariaceae Genus: Pinguicula L. Species[1] 126, see separate list. Synonyms[1] Brandonia Rchb. Isoloba Raf. Pinguicola Zumagl.

***Pinguicula***, commonly known as **butterworts**, is a genus of [carnivorous](/source/Carnivorous_plant) [flowering plants](/source/Flowering_plant) in the family [Lentibulariaceae](/source/Lentibulariaceae). They use sticky, glandular leaves to lure, trap, and digest insects in order to supplement the poor mineral nutrition they obtain from the environment. 126 species are currently accepted.[1] 13 are native to Europe, 9 to North America, and some to northern Asia. The largest number of species is in South and Central America.

## Etymology

The name *Pinguicula* is derived from a term coined by [Conrad Gesner](/source/Conrad_Gesner), who in his 1561 work entitled *Horti Germaniae* commented on the glistening leaves: *"propter pinguia et tenera folia…"* (Latin *pinguis*, "fat"). The common name "butterwort" reflects this characteristic.[2]

## Characteristics

The majority of *Pinguicula* are [perennial plants](/source/Perennial_plant). The only known [annuals](/source/Annual_plant) are [*P. sharpii*](https://en.wikipedia.org/w/index.php?title=Pinguicula_sharpii&action=edit&redlink=1), [*P. takakii*](https://en.wikipedia.org/w/index.php?title=Pinguicula_takakii&action=edit&redlink=1), [*P. crenatiloba*](https://en.wikipedia.org/w/index.php?title=Pinguicula_crenatiloba&action=edit&redlink=1), and [*P. pumila*](/source/Pinguicula_pumila). All species form stemless [rosettes](/source/Rosette_(botany)).

### Habitat

Winter rosette of *[Pinguicula cyclosecta](/source/Pinguicula_cyclosecta)* (non-carnivorous phase)

Summer rosette of *[Pinguicula cyclosecta](/source/Pinguicula_cyclosecta)* (carnivorous phase)

Butterworts can be divided roughly into two main groups based on the climate in which they grow; each group is then further subdivided based on [morphological](/source/Morphology_(biology)) characteristics. Although these groups are not [cladistically](/source/Cladistics) supported by genetic studies,[3] these groupings are nonetheless convenient for horticultural purposes.

Tropical butterworts form somewhat compact winter rosettes composed of fleshy leaves or retain carnivorous leaves year-round.[4] They are typically located in regions where water is least seasonally plentiful, as too damp soil conditions can lead to rotting. They are found in areas in which nitrogenous resources are known to be in low levels, infrequent or unavailable, due to acidic soil conditions.

[Temperate](/source/Temperate) species often form tight buds (called [hibernacula](/source/Hibernaculum_(botany))) composed of scale-like leaves during a winter [dormancy](/source/Dormancy) period. During this time the roots (with the exception of [*P. alpina*](/source/Pinguicula_alpina)) and carnivorous leaves wither.[5] Temperate species flower when they form their summer rosettes while tropical species flower at each rosette change.

Many butterworts cycle between rosettes composed of carnivorous and non-carnivorous leaves as the seasons change, so these two ecological groupings can be further divided according to their ability to produce different leaves during their growing season. If the growth in the summer is different in size or shape to that in the early spring (for temperate species) or in the winter (tropical species), then plants are considered [heterophyllous](/source/Heterophyllous); whereas uniform growth identifies a [homophyllous](https://en.wikipedia.org/w/index.php?title=Homophyllous&action=edit&redlink=1) species.

This results in four groupings:

- **Tropical butterworts**: species which do not undergo a winter dormancy but continue to alternately bloom and form rosettes. - ***Heterophyllous tropical species***: species that alternate between rosettes of carnivorous leaves during the warm season and compact rosettes of fleshy non-carnivorous leaves during the cool season. Examples include [*P. moranensis*](/source/Pinguicula_moranensis), [*P. gypsicola*](/source/Pinguicula_gypsicola), and [*P. laxifolia*](/source/Pinguicula_laxifolia). - ***Homophyllous tropical species***: these species produce rosettes of carnivorous leaves of roughly uniform size throughout the year, such as [*P. gigantea*](/source/Pinguicula_gigantea).

- **Temperate butterworts**: these plants are native to climate zones with cold winters. They produce a winter-resting bud ([hibernaculum](/source/Hibernaculum_(botany))) during the winter. - ***Heterophyllous temperate species***: species where the vegetative and generative rosettes differ in shape and/or size, as seen in [*P. lutea*](/source/Pinguicula_lutea) and [*P. lusitanica*](/source/Pinguicula_lusitanica). - ***Homophyllous temperate species***: the vegetative and generative rosettes appear identical, as exhibited by [*P. alpina*](/source/Pinguicula_alpina), [*P. grandiflora*](/source/Pinguicula_grandiflora), and [*P. vulgaris*](/source/Pinguicula_vulgaris).

### Roots

The [root system](/source/Root) of *Pinguicula* species is relatively undeveloped. The thin, white roots serve mainly as an anchor for the plant and to absorb moisture (nutrients are absorbed through carnivory). In temperate species these roots wither (except in [*P. alpina*](/source/Pinguicula_alpina)) when the [hibernaculum](/source/Hibernaculum_(botany)) is formed. In the few [epiphytic](/source/Epiphytic) species (such as [*P. lignicola*](https://en.wikipedia.org/w/index.php?title=Pinguicula_lignicola&action=edit&redlink=1)), the roots form anchoring suction cups.

### Leaves and carnivory

A fly trapped on a butterwort leaf. Glandular hairs are visible.

The leaf blade of a butterwort is smooth, rigid, and succulent, usually bright green or pinkish in colour. Depending on species, the leaves are between 2 and 30 cm (1–12") long. The leaf shape depends on the species, but is usually roughly [obovate](/source/Leaf#Terminology), [spatulate](/source/Leaf#Terminology), or [linear](/source/Leaf#Terminology).[6] They can also appear yellow in color with a soft feel and a greasy consistency to the leaves.

Vector graphic of the trapping and digestive features of a *Pinguicula* leaf

Like all members of the family [Lentibulariaceae](/source/Lentibulariaceae), butterworts are carnivorous.[7] The mechanistic actions that these plants use to lure and capture prey is through a means of sticky or adhesives substances that are produced by mucilage secreted by glands located on the leaf's surface. In order to catch and digest insects, the leaf of a butterwort uses two specialized glands which are scattered across the leaf surface (usually only on the upper surface, with the exception of [*P. gigantea*](/source/Pinguicula_gigantea) and *[P. longifolia ssp. longifolia](/source/Pinguicula_longifolia)*).[5]

Pinguicula esseriana in the greenhouse of the Kharkiv Botanical Garden

One is termed a *peduncular gland*, and consists of a few [secretory](/source/Secrete) cells on top of a single stalk cell. These cells produce a [mucilaginous](/source/Mucilage) secretion which forms visible droplets across the leaf surface. This wet appearance probably helps lure prey in search of water (a similar phenomenon is observed in the [sundews](/source/Drosera)). The droplets secrete limited amounts of digestive enzymes, and serve mainly to entrap insects. On contact with an insect, the peduncular glands release additional mucilage from special reservoir cells located at the base of their stalks.[5] The insect will begin to struggle, triggering more glands and encasing itself in mucilage. Some species can bend their leaf edges slightly by [thigmotropism](/source/Thigmotropism), bringing additional glands into contact with the trapped insect.[5]

The second type of gland found on butterwort leaves are *sessile glands* which lie flat on the leaf surface. Once the prey is entrapped by the peduncular glands and digestion begins, the initial flow of nitrogen triggers enzyme release by the sessile glands.[5] These enzymes, which include [amylase](/source/Amylase), [esterase](/source/Esterase), [phosphatase](/source/Phosphatase), [protease](/source/Protease), and [ribonuclease](/source/Ribonuclease) break down the digestible components of the insect body. These fluids are then absorbed back into the leaf surface through [cuticular](/source/Plant_cuticle) holes, leaving only the [chitin](/source/Chitin) [exoskeleton](/source/Exoskeleton) of the larger insects on the leaf surface.

The holes in the cuticle which allow for this digestive mechanism also pose a challenge for the plant, since they serve as breaks in the [cuticle](/source/Plant_cuticle) (waxy layer) that protects the plant from [desiccation](/source/Desiccation). As a result, most butterworts live in humid environments.

Flower of *P. vulgaris*

Butterworts are usually only able to trap small insects and those with large wing surfaces. They can also digest [pollen](/source/Pollen) which lands on their leaf surface. The secretory system can only function a single time, so that a particular area of the leaf surface can only be used to digest insects once.[5]

Unlike many other carnivorous plant species, butterworts do not appear to use [jasmonates](/source/Jasmonates) as a control system to switch on the production of digestive enzymes. Jasmonates are involved in the butterwort's defense against attacking insects, but not in its response to prey.[8][9][10] Of the eight enzymes identified in the digestive secretions of butterworts, [alpha-amylase](/source/Alpha-amylase) appears to be unique when compared to other carnivorous plants. This research suggests that butterwort may have co-opted a different set of genes in its development of carnivory.[10]

### Flowers

The flower of a hybrid butterwort

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As with almost all [carnivorous plants](/source/Carnivorous_plants), the flowers of butterworts are held far above the rest of the plant by a long stalk, in order to reduce the probability of trapping potential pollinators.[11] The single, long-lasting flowers are [zygomorphic](/source/Zygomorphic), with two lower lip petals characteristic of the [bladderwort family](/source/Lentibulariaceae), and a spur extending from the back of the flower. The [calyx](/source/Sepals) has five sepals, and the petals are arranged in a two-part lower lip and a three-part upper lip.[12] Most butterwort flowers are blue, violet or white, often suffused with a yellow, greenish or reddish tint. *[P. laueana](/source/Pinguicula_laueana)* and the newly described [*P. caryophyllacea*](/source/Pinguicula_caryophyllacea) are unique in having a strikingly red flowers. Butterworts are often cultivated and hybridized primarily for their flowers.

The shape and colors of butterwort flowers are distinguishing characteristics which are used to divide the genus into subgenera and to distinguish individual species from one another.[13]

### Fruit and seed

The round to egg-shaped [seed capsules](/source/Capsule_(fruit)) open when dry into two halves, exposing numerous small (0.5–1 mm), brown seeds. If moisture is present the [silique](/source/Silique) closes, protecting the seed and opening again upon dryness to allow for [wind dispersal](/source/Seed_dispersal#Wind). Many species have a net-like pattern on their seed surface to allow them to land on water surfaces without sinking, since many non-epiphytic butterworts grow near water sources. The [haploid](/source/Haploid) [chromosome](/source/Chromosome) number of butterworts is either n = 8 or n = 11 (or a multiple thereof), depending on species. The exception is *[P. lusitanica](/source/Pinguicula_lusitanica)*, whose chromosome count is n = 6.[*[citation needed](https://en.wikipedia.org/wiki/Wikipedia:Citation_needed)*]

### Diet

The diet will range depending on the taxonomy and size of the prey due to the plant's retention ability. These size limitations are known to be the main element influencing what prey sources this carnivorous plant can access.[14] They can also acquire nourishment from pollen and other plant parts that are high in protein, as other plants can become trapped on their leaves, thus, butterworts are both carnivorous and herbivorous plants.[7] The diet consists of several species from the arthropod taxa; the majority of their prey are insects that have wings and are able to fly. The luring, retaining, and seizing of prey is the first steps in the feeding procedure for carnivorous plants; the result of the process is absorption and digestion of nutrients sourced from these food supplies. Pinguicula species do not select their prey, as they passively accumulate them through methods of sticky, adhesive leaves. However, they do have the ability of visual attraction of their colorful leaves, which will increase the likelihood of luring and capturing a specific taxa.[15] [*[better source needed](https://en.wikipedia.org/wiki/Wikipedia:Verifiability#Questionable_sources)*] Pinguicula capture their food source/ prey by means of the mucilaginous, sticky substances produced by their stalk glands on the top of their leaf. Once the prey has become trapped in the peduncular glands, the sessile glands present will then produce enzymes needed to accomplish digestion and breaking down the digestible regions of the prey for their nutrients; taking in the fluids of the food source by means of cuticular holes present on the leaf's surface.

### Vegetative propagation

As well as [sexual reproduction](/source/Sexual_reproduction) by seed, many butterworts can reproduce [asexually](/source/Asexual_reproduction) by [vegetative reproduction](/source/Vegetative_reproduction). Many members of the genus form offshoots during or shortly after flowering (*e.g.*, *[P. vulgaris](/source/Pinguicula_vulgaris)*), which grow into new [genetically identical](/source/Cloning) adults. A few other species form new [offshoots](/source/Offshoot_(plant)) using [stolons](/source/Stolons) (*e.g.*, *[P. calyptrata](https://en.wikipedia.org/w/index.php?title=Pinguicula_calyptrata&action=edit&redlink=1)*, *[P. vallisneriifolia](https://en.wikipedia.org/w/index.php?title=Pinguicula_vallisneriifolia&action=edit&redlink=1)*) while others form plantlets at the leaf margins (*e.g.*, *[P. heterophylla](/source/Pinguicula_heterophylla)*, *[P. primuliflora](/source/Pinguicula_primuliflora)*).

## Distribution

*Pinguicula* distribution

Butterworts are distributed throughout the [Northern Hemisphere](/source/Northern_Hemisphere) ([map](http://www.pinguicula.org/pages/culture/Overview_fichiers/image002.jpg)). The greatest concentration of species, however, is in humid mountainous regions of Mexico, Central America and South America, where populations can be found as far south as [Tierra del Fuego](/source/Tierra_del_Fuego). Australia and Antarctica are the only continents without any native butterworts.

While the highest diversity of species is observed in Central America (roughly 50% of butterwort species are found here), the most ancestral species of the genus *Pinguicula*, both in terms of morphology and molecular reconstruction, are the group of temperate homophyllous species (examples include *P. lusitanica*, *P. hirtiflora*, *[P. antartica](https://en.wikipedia.org/w/index.php?title=Pinguicula_antartica&action=edit&redlink=1)*, *[P. lutea](/source/Pinguicula_lutea)*, etc.). Likewise, as other members of [Lentibulariaceae](/source/Lentibulariaceae) family, *Pinguicula* ancestors evolved in South America, where Andean species most probably present as remnants of the ancestral stock of South American *Pinguicula* adapted to highland conditions with the rise of the Andes [16][17][18]

The great majority of individual *Pinguicula* species have a very limited [distribution](/source/Population_distribution), often restricted to few canyons or mountain cliffs. The two butterwort species with the widest distribution - *[P. alpina](/source/Pinguicula_alpina)* and *[P. vulgaris](/source/Pinguicula_vulgaris)* - are found throughout much of Europe and North America and in some regions in Asia. Other species found in North America include *[P. caerulea](/source/Pinguicula_caerulea)*, *[P. ionantha](/source/Pinguicula_ionantha)*, *[P. lutea](/source/Pinguicula_lutea)*, *[P. macroceras](/source/Pinguicula_macroceras)*, *[P. planifolia](/source/Pinguicula_planifolia)*, *[P. primuliflora](/source/Pinguicula_primuliflora)*, *[P. pumila](/source/Pinguicula_pumila)*, and *[P. villosa](https://en.wikipedia.org/w/index.php?title=Pinguicula_villosa&action=edit&redlink=1)*.[*[citation needed](https://en.wikipedia.org/wiki/Wikipedia:Citation_needed)*]

## Habitat

*P. macroceras ssp. nortensis* growing on a wet rock wall in northern California.

*P. leptoceras* in alpine grassland in Südtirol, Italy

In general, butterworts grow in nutrient-poor, alkaline soils. Some species have adapted to other soil types, such as acidic [peat bogs](/source/Peat_bog) (ex. *[P. vulgaris](/source/Pinguicula_vulgaris)*, *[P. calyptrata](https://en.wikipedia.org/w/index.php?title=Pinguicula_calyptrata&action=edit&redlink=1)*, *[P. lusitanica](/source/Pinguicula_lusitanica)*), soils composed of pure [gypsum](/source/Gypsum) (*[P. gypsicola](/source/Pinguicula_gypsicola)* and other Mexican species), or even vertical rock walls (*[P. ramosa](/source/Pinguicula_ramosa)*, *[P. vallisneriifolia](https://en.wikipedia.org/w/index.php?title=Pinguicula_vallisneriifolia&action=edit&redlink=1)*, and most of the Mexican species). A few species are epiphytes (*[P. casabitoana](/source/Pinguicula_casabitoana)*, *[P. hemiepiphytica](/source/Pinguicula_hemiepiphytica)*, *[P. lignicola](https://en.wikipedia.org/w/index.php?title=Pinguicula_lignicola&action=edit&redlink=1)*). Many of the Mexican species commonly grow on mossy banks, rock, and roadsides in oak-pine forests. *Pinguicula macroceras ssp. nortensis* has even been observed growing on hanging dead grasses. *[P. lutea](/source/Pinguicula_lutea)* grows in pine [flatwoods](/source/Flatwoods).[19] Other species, such as *[P. vulgaris](/source/Pinguicula_vulgaris)*, grow in [fens](/source/Fen). Secondary habitats, such as [anthropogenic biomes](/source/Anthropogenic_biome), might include steep rock walls along the roads[20] or roadside ditches[21]. Each of these environments is nutrient-poor, allowing butterworts to escape competition from other canopy-forming species, particularly grasses and sedges.[22]

Butterworts need habitats that are almost constantly moist or wet, at least during their carnivorous growth stage. Many Mexican species lose their carnivorous leaves, and sprout succulent leaves, or die back to onion-like "bulbs" to survive the winter drought, at which point they can survive in bone-dry conditions. The moisture they need for growing can be supplied by either a high [groundwater table](/source/Water_table), or by high humidity or high precipitation. Unlike many other carnivorous plants that require sunny locations, many butterworts thrive in part-sun or even shady conditions.

## Conservation status

The environmental threats faced by various *Pinguicula* species depend on their location and on how widespread their distribution is. Most endangered are the species which are [endemic](/source/Endemic_(ecology)) to small areas, such as *[P. ramosa](/source/Pinguicula_ramosa)*, *[P. casabitoana](/source/Pinguicula_casabitoana)*, and *[P. fiorii](https://en.wikipedia.org/w/index.php?title=Pinguicula_fiorii&action=edit&redlink=1)*. These populations are threatened primarily by [habitat destruction](/source/Habitat_destruction). [Wetland](/source/Wetland) destruction has threatened several US species. Most of these are federally listed as either threatened or endangered, and *[P. ionantha](/source/Pinguicula_ionantha)* is listed on [CITES](/source/CITES) appendix I, giving it additional protection.[*[citation needed](https://en.wikipedia.org/wiki/Wikipedia:Citation_needed)*]

## Botanical history

*Pinguicula vulgaris*, illustration

The first mention of butterworts in botanical literature is an entry entitled *Zitroch chrawt oder schmalz chrawt* ("lard herb") by [Vitus Auslasser](https://en.wikipedia.org/w/index.php?title=Vitus_Auslasser&action=edit&redlink=1) in his 1479 work on medicinal herbs entitled *Macer de viribus Herbarum*.[23] The name *Zittrochkraut* is still used for butterworts in [Tirol](/source/Tyrol_(state)), Austria.

In 1583, [Clusius](/source/Carolus_Clusius) already distinguished between two forms in his *Historia stirpium rariorum per Pannoniam, Austriam*: a blue-flowered form (*[P. vulgaris](/source/Pinguicula_vulgaris)*) and a white-flowered form (*[Pinguicula alpina](/source/Pinguicula_alpina)*). [Linnaeus](/source/Carl_Linnaeus) added *[P. villosa](https://en.wikipedia.org/w/index.php?title=Pinguicula_villosa&action=edit&redlink=1)* and *[P. lusitanica](/source/Pinguicula_lusitanica)* when he published his *[Species Plantarum](/source/Species_Plantarum)* in 1753. The number of known species rose sharply with the exploration of the new continents in the 19th century; by 1844, 32 species were known.

It was only in the late 19th century that the carnivory of this genus began to be studied in detail. In a letter to [Asa Gray](/source/Asa_Gray) dated June 3, 1874, [Charles Darwin](/source/Charles_Darwin) mentioned his early observations of the butterwort's digestive process and [insectivorous](/source/Insectivorous) nature.[24] Darwin studied these plants extensively.[25] [S. J. Casper's](/source/Siegfried_Jost_Casper) large 1966 monograph of the genus[26] included 46 species, a number which has almost doubled since then. Many exciting discoveries have been made in recent years, especially in Mexico. Another important development in the history of butterworts is the formation of the [International Pinguicula Study Group](/source/International_Pinguicula_Study_Group), an organization dedicated to furthering the knowledge of this genus and promoting its popularity in cultivation, in the 1990s.

## Uses

Butterworts are widely cultivated by carnivorous plant enthusiasts. The temperate species and many of the Mexican butterworts are relatively easy to grow and have therefore gained relative popularity. Two of the most widely grown plants are the hybrid [cultivars](/source/Cultivar) *Pinguicula* × 'Sethos' and *Pinguicula* × 'Weser'. Both are crosses of *[Pinguicula ehlersiae](https://en.wikipedia.org/w/index.php?title=Pinguicula_ehlersiae&action=edit&redlink=1)* and *[Pinguicula moranensis](/source/Pinguicula_moranensis)*, and are employed by commercial [orchid](/source/Orchid) nurseries to combat pests.[*[citation needed](https://en.wikipedia.org/wiki/Wikipedia:Citation_needed)*]

Butterworts also produce a strong bactericide which prevents insects from rotting while they are being digested. According to [Linnaeus](/source/Carl_Linnaeus), this property has long been known by northern Europeans, who applied butterwort leaves to the [sores](/source/Ulcer_(dermatology)) of cattle to promote healing.[27] Additionally, butterwort leaves were used to [curdle](/source/Curdle) milk and form a buttermilk-like [fermented milk product](/source/Fermented_milk_product) called *[filmjölk](/source/Filmj%C3%B6lk)* (Sweden) and *tjukkmjølk* (Norway).[28]

## Classification

See also: [List of Pinguicula species](/source/List_of_Pinguicula_species)

*Pinguicula* belong to the bladderwort family ([Lentibulariaceae](/source/Lentibulariaceae)), along with *[Utricularia](/source/Utricularia)* and *[Genlisea](/source/Genlisea)*. [Siegfried Jost Casper](/source/Siegfried_Jost_Casper) [systematically](/source/Systematics) divided them into three [subgenera](/source/Subgenus) with 15 [sections](/source/Section_(botany)).[26]

A detailed study of the phylogenetics of butterworts by Cieslak *et al.* (2005)[3] found that all of the currently accepted [subgenera](/source/Subgenus) and many of the [sections](/source/Section_(biology)) were [polyphyletic](/source/Polyphyletic). The diagram below gives a more accurate representation of the correct [cladogram](/source/Cladogram). Polyphyletic sections are marked with an *****.

                  ┌────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)

## References

1. ^ [***a***](#cite_ref-powo_1-0) [***b***](#cite_ref-powo_1-1) [***c***](#cite_ref-powo_1-2) ["*Pinguicula* L."](https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30014019-2) *[Plants of the World Online](/source/Plants_of_the_World_Online)*. [Royal Botanic Gardens, Kew](/source/Royal_Botanic_Gardens%2C_Kew). Retrieved 26 April 2025.

1. **[^](#cite_ref-2)** ["How to Grow Your Indoor Butterworts (Pinguicula)"](https://www.ukhouseplants.com/plants/pinguicula-butterworts). *UKHouseplants*. Retrieved 11 March 2022.

1. ^ [***a***](#cite_ref-cieslak2005_3-0) [***b***](#cite_ref-cieslak2005_3-1) Cieslak T, Polepalli JS, White A, Müller K, Borsch T, Barthlott W, Steiger J, Marchant A, Legendre L (2005). ["Phylogenetic analysis of *Pinguicula* (Lentibulariaceae): chloroplast DNA sequences and morphology support several geographically distinct radiations"](https://doi.org/10.3732%2Fajb.92.10.1723). *[American Journal of Botany](/source/American_Journal_of_Botany)*. **92** (10): 1723–1736. [Bibcode](/source/Bibcode_(identifier)):[2005AmJB...92.1723C](https://ui.adsabs.harvard.edu/abs/2005AmJB...92.1723C). [doi](/source/Doi_(identifier)):[10.3732/ajb.92.10.1723](https://doi.org/10.3732%2Fajb.92.10.1723). [PMID](/source/PMID_(identifier)) [21646090](https://pubmed.ncbi.nlm.nih.gov/21646090).

1. **[^](#cite_ref-4)** ["Carnivorous plant | botany"](https://www.britannica.com/plant/carnivorous-plant). *Encyclopedia Britannica*. Retrieved 2020-03-01.

1. ^ [***a***](#cite_ref-legendre2000_5-0) [***b***](#cite_ref-legendre2000_5-1) [***c***](#cite_ref-legendre2000_5-2) [***d***](#cite_ref-legendre2000_5-3) [***e***](#cite_ref-legendre2000_5-4) [***f***](#cite_ref-legendre2000_5-5) Legendre L (2002). ["The genus *Pinguicula* L. (Lentibulariaceae): an overview"](http://www.pinguicula.org/pages/culture/Overview.htm). *Acta Botanica Gallica*. **141** (1): 77–95.

1. **[^](#cite_ref-6)** ["Carnivorous Butterwort Care – How To Grow Butterworts"](https://www.gardeningknowhow.com/houseplants/butterwort/how-to-grow-butterworts.htm). *Gardening Know How*. 25 March 2013. Retrieved 2020-03-01.

1. ^ [***a***](#cite_ref-pmid28313028_7-0) [***b***](#cite_ref-pmid28313028_7-1) Zamora, R. (1990). "The feeding ecology of a carnivorous plant (Pinguicula nevadense): Prey analysis and capture constraints". *Oecologia*. **84** (3): 376–379. [Bibcode](/source/Bibcode_(identifier)):[1990Oecol..84..376Z](https://ui.adsabs.harvard.edu/abs/1990Oecol..84..376Z). [doi](/source/Doi_(identifier)):[10.1007/bf00329762](https://doi.org/10.1007%2Fbf00329762). [PMID](/source/PMID_(identifier)) [28313028](https://pubmed.ncbi.nlm.nih.gov/28313028). [S2CID](/source/S2CID_(identifier)) [8038140](https://api.semanticscholar.org/CorpusID:8038140).

1. **[^](#cite_ref-Pain_8-0)** Pain, Stephanie (2 March 2022). ["How plants turned predator"](https://knowablemagazine.org/article/living-world/2022/how-plants-turned-predator). *Knowable Magazine*. [doi](/source/Doi_(identifier)):[10.1146/knowable-030122-1](https://doi.org/10.1146%2Fknowable-030122-1). Retrieved 11 March 2022.

1. **[^](#cite_ref-Hedrich_9-0)** Hedrich, Rainer; Fukushima, Kenji (17 June 2021). ["On the Origin of Carnivory: Molecular Physiology and Evolution of Plants on an Animal Diet"](https://doi.org/10.1146%2Fannurev-arplant-080620-010429). *Annual Review of Plant Biology*. **72** (1): 133–153. [Bibcode](/source/Bibcode_(identifier)):[2021AnRPB..72..133H](https://ui.adsabs.harvard.edu/abs/2021AnRPB..72..133H). [doi](/source/Doi_(identifier)):[10.1146/annurev-arplant-080620-010429](https://doi.org/10.1146%2Fannurev-arplant-080620-010429). [ISSN](/source/ISSN_(identifier)) [1543-5008](https://search.worldcat.org/issn/1543-5008). [PMID](/source/PMID_(identifier)) [33434053](https://pubmed.ncbi.nlm.nih.gov/33434053). [S2CID](/source/S2CID_(identifier)) [231595236](https://api.semanticscholar.org/CorpusID:231595236).

1. ^ [***a***](#cite_ref-Kocáb_10-0) [***b***](#cite_ref-Kocáb_10-1) Kocáb, Ondřej; Jakšová, Jana; Novák, Ondřej; Petřík, Ivan; Lenobel, René; Chamrád, Ivo; Pavlovič, Andrej (22 June 2020). ["Jasmonate-independent regulation of digestive enzyme activity in the carnivorous butterwort Pinguicula × Tina"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7307851). *Journal of Experimental Botany*. **71** (12): 3749–3758. [doi](/source/Doi_(identifier)):[10.1093/jxb/eraa159](https://doi.org/10.1093%2Fjxb%2Feraa159). [PMC](/source/PMC_(identifier)) [7307851](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7307851). [PMID](/source/PMID_(identifier)) [32219314](https://pubmed.ncbi.nlm.nih.gov/32219314).

1. **[^](#cite_ref-11)** ["Common Butterwort (Pinguicula vulgaris)"](https://easyscape.com/species/Pinguicula-vulgaris%28Common-Butterwort%29). *easyscape.com*. Retrieved 2026-01-20.

1. **[^](#cite_ref-12)** Lustofin, Krzysztof; Świątek, Piotr; Miranda, Vitor F. O.; Płachno, Bartosz J. (2020-01-01). ["Flower nectar trichome structure of carnivorous plants from the genus butterworts Pinguicula L. (Lentibulariaceae)"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6982637). *Protoplasma*. **257** (1): 245–259. [Bibcode](/source/Bibcode_(identifier)):[2020Prpls.257..245L](https://ui.adsabs.harvard.edu/abs/2020Prpls.257..245L). [doi](/source/Doi_(identifier)):[10.1007/s00709-019-01433-8](https://doi.org/10.1007%2Fs00709-019-01433-8). [ISSN](/source/ISSN_(identifier)) [1615-6102](https://search.worldcat.org/issn/1615-6102). [PMC](/source/PMC_(identifier)) [6982637](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6982637). [PMID](/source/PMID_(identifier)) [31428856](https://pubmed.ncbi.nlm.nih.gov/31428856).

1. **[^](#cite_ref-13)** ["On the infrageneric classification of Pinguicula"](https://cpn.carnivorousplants.org/articles/CPNv50n4p174_188.pdf) (PDF).

1. **[^](#cite_ref-14)** Zamora, Regino (1990). "The Feeding Ecology of a Carnivorous Plant (Pinguicula nevadense): Prey Analysis and Capture Constraints". *Oecologia*. **84** (3): 376–379. [Bibcode](/source/Bibcode_(identifier)):[1990Oecol..84..376Z](https://ui.adsabs.harvard.edu/abs/1990Oecol..84..376Z). [doi](/source/Doi_(identifier)):[10.1007/BF00329762](https://doi.org/10.1007%2FBF00329762). [ISSN](/source/ISSN_(identifier)) [0029-8549](https://search.worldcat.org/issn/0029-8549). [JSTOR](/source/JSTOR_(identifier)) [4219437](https://www.jstor.org/stable/4219437). [PMID](/source/PMID_(identifier)) [28313028](https://pubmed.ncbi.nlm.nih.gov/28313028). [S2CID](/source/S2CID_(identifier)) [8038140](https://api.semanticscholar.org/CorpusID:8038140).

1. **[^](#cite_ref-15)** ["All About the Butterworts Plant"](https://www.carnivorous--plants.com/butterworts-plant.html). *www.carnivorous--plants.com*. Retrieved 2020-03-16.

1. **[^](#cite_ref-16)** Brittnacher, John (2010). "Evolution of the Lamiales Carnivores". International Carnivorous Plant Society. {{[cite web](https://en.wikipedia.org/wiki/Template:Cite_web)}}: Missing or empty |url= ([help](https://en.wikipedia.org/wiki/Help:CS1_errors#cite_web_url))

1. **[^](#cite_ref-17)** Lampard, Stan; Gluch, Oliver; Robinson, Alastair S.; Fleischmann, Andreas; Temple, Paul; McPherson, Stewart; Roccia, Aymeric; Partrat, Eric; Legendre, Laurent (2016). *Pinguicula of Latin America*. Redfern Natural History. [ISBN](/source/ISBN_(identifier)) [978-1-908787-15-6](https://en.wikipedia.org/wiki/Special:BookSources/978-1-908787-15-6).

1. **[^](#cite_ref-18)** Shimai, H.; Setoguchi, H.; Roberts, D. L.; Sun, M. (2021). ["Biogeographical patterns and speciation of the genus Pinguicula (Lentibulariaceae) inferred by phylogenetic analyses"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8184156). *PLOS ONE*. **16** (6) e0252581. [Bibcode](/source/Bibcode_(identifier)):[2021PLoSO..1652581S](https://ui.adsabs.harvard.edu/abs/2021PLoSO..1652581S). [doi](/source/Doi_(identifier)):[10.1371/journal.pone.0252581](https://doi.org/10.1371%2Fjournal.pone.0252581). [PMC](/source/PMC_(identifier)) [8184156](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8184156). [PMID](/source/PMID_(identifier)) [34097720](https://pubmed.ncbi.nlm.nih.gov/34097720).

1. **[^](#cite_ref-19)** Keddy PA, Smith L, Campbell DR, Clark M, Montz G (2006). "Patterns of herbaceous plant diversity in southeastern Louisiana pine savannas". *Applied Vegetation Science*. **9** (1): 17–26. [Bibcode](/source/Bibcode_(identifier)):[2006AppVS...9...17K](https://ui.adsabs.harvard.edu/abs/2006AppVS...9...17K). [doi](/source/Doi_(identifier)):[10.1111/j.1654-109X.2006.tb00652.x](https://doi.org/10.1111%2Fj.1654-109X.2006.tb00652.x).

1. **[^](#cite_ref-20)** ["Pinguicula acuminata"](https://www.pinguicula.org/plants/acuminata/). *Pinguicula.org*. 18 January 2026. Retrieved 2026-04-19.

1. **[^](#cite_ref-21)** ["Pinguicula ionantha (Violet-flowered Butterwort)"](https://saveplants.org/plant-profile/3460/Pinguicula-ionantha/Violet-flowered-Butterwort/). *Center for Plant Conservation*. Retrieved 2026-04-19.

1. **[^](#cite_ref-22)** Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. Chapter 5.

1. **[^](#cite_ref-auslasser-1479_23-0)** Auslasser, Vitus (1479). [*Macer de viribus herbarum*](https://www.digitale-sammlungen.de/en/view/bsb00092488?page=547). Digitised by Bayerische Staatsbibliothek, München. p. 272r. Retrieved 6 April 2026.

1. **[^](#cite_ref-24)** Darwin, Charles (2015). [*The Correspondence of Charles Darwin: Volume 22, 1874*](https://books.google.com/books?id=_AK7BwAAQBAJ&q=%22Day+before+Yesterday+I+found+out+that+Pinguicula+digests%22). [Cambridge University Press](/source/Cambridge_University_Press). p. 487. [ISBN](/source/ISBN_(identifier)) [978-1-316-24095-3](https://en.wikipedia.org/wiki/Special:BookSources/978-1-316-24095-3).

1. **[^](#cite_ref-25)** [Darwin](/source/Charles_Darwin) C (1875). [*Insectivorous plants*](https://web.archive.org/web/20060923021622/http://pages.britishlibrary.net/charles.darwin3/insectivorous/insect01.htm). London: John Murray. [ISBN](/source/ISBN_(identifier)) [1-4102-0174-0](https://en.wikipedia.org/wiki/Special:BookSources/1-4102-0174-0). Archived from [the original](http://pages.britishlibrary.net/charles.darwin3/insectivorous/insect01.htm) on 2006-09-23. {{[cite book](https://en.wikipedia.org/wiki/Template:Cite_book)}}: ISBN / Date incompatibility ([help](https://en.wikipedia.org/wiki/Help:CS1_errors#invalid_isbn_date))

1. ^ [***a***](#cite_ref-casper1966_26-0) [***b***](#cite_ref-casper1966_26-1) Casper SJ (1966). Monographie der Gattung *Pinguicula* L.*(Heft 127/128, Vol 31)*. Stuttgart: [Bibliotheca Botanica](/source/Bibliotheca_Botanica).

1. **[^](#cite_ref-27)** D'Amato P (1988). [*The Savage Garden: Cultivating Carnivorous Plants*](/source/The_Savage_Garden%3A_Cultivating_Carnivorous_Plants). Ten Speed Press. [ISBN](/source/ISBN_(identifier)) [0-89815-915-6](https://en.wikipedia.org/wiki/Special:BookSources/0-89815-915-6).

1. **[^](#cite_ref-28)** Amilien, Virginie; Torjusen, Hanne; Vittersø, Gunnar (2005). ["From local food to terroir product ? - Some views about Tjukkmjølk, the traditional thick sour milk from Røros, Norway"](https://doi.org/10.4000%2Faof.211). *Anthropology of Food*. **4** (4). [doi](/source/Doi_(identifier)):[10.4000/aof.211](https://doi.org/10.4000%2Faof.211).

## Further reading

- Barthlott W, Porembski S, Seine R, Theisen I (2004). *Karnivoren*. Stuttgart: Verlag Eugen Ulmer. [ISBN](/source/ISBN_(identifier)) [3-8001-4144-2](https://en.wikipedia.org/wiki/Special:BookSources/3-8001-4144-2).

- Müller K, Borsch T, Legendre L, Porembski S, Theisen I, Barthlott W (2004). "Evolution of carnivory in Lamiales". *Plant Biology*. **6** (4): 1–14. [Bibcode](/source/Bibcode_(identifier)):[2004PlBio...6..477M](https://ui.adsabs.harvard.edu/abs/2004PlBio...6..477M). [doi](/source/Doi_(identifier)):[10.1055/s-2004-817909](https://doi.org/10.1055%2Fs-2004-817909). [PMID](/source/PMID_(identifier)) [15248131](https://pubmed.ncbi.nlm.nih.gov/15248131).

- Keddy, P.A. (2010). Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK.

- Givnish, T. J. (1988). Ecology and evolution of carnivorous plants. In Plant–Animal Interactions, ed. W. B. Abrahamson, pp. 243–90. New York: McGraw-Hill.

## External links

- Media related to [Pinguicula](https://commons.wikimedia.org/wiki/Pinguicula) at Wikimedia Commons

- Data related to [Pinguicula](https://species.wikimedia.org/wiki/Pinguicula) at Wikispecies

- [An exhaustive website on the genus *Pinguicula*](http://www.pinguicula.org/)

- Schlauer, J. [Carnivorous Plant Database](http://www.omnisterra.com/bot/cp_home.cgi) [Archived](https://web.archive.org/web/20160918161537/http://www.omnisterra.com/bot/cp_home.cgi) 2016-09-18 at the [Wayback Machine](/source/Wayback_Machine), version 15 November 16: 25.

- [Flora Europaea: *Pinguicula* species list](http://rbg-web2.rbge.org.uk/cgi-bin/nph-readbtree.pl/feout?FAMILY_XREF=&GENUS_XREF=Pinguicula&SPECIES_XREF=&TAXON_NAME_XREF=&RANK=species)

- [Botanical Society of America, *Pinguicula* - the Butterworts](http://www.botany.org/Carnivorous_Plants/Pinguicula.php) [Archived](https://web.archive.org/web/20130103153551/http://www.botany.org/Carnivorous_Plants/Pinguicula.php) 2013-01-03 at the [Wayback Machine](/source/Wayback_Machine)

v t e Carnivorous and protocarnivorous plants Carnivorous genera († extinct) Aldrovanda †Archaeamphora? Brocchinia Byblis Catopsis Cephalotus Darlingtonia Dionaea Drosera †Droserapites †Droserapollis †Droseridites Drosophyllum †Fischeripollis Genlisea Heliamphora Nepenthes †Palaeoaldrovanda? Philcoxia Pinguicula Roridula Sarracenia †Saxonipollis Triantha Triphyophyllum Utricularia Protocarnivorous genera Aracamunia Capsella Colura Dipsacus Drymocallis Geranium Ibicella Lathraea Paepalanthus Passiflora Plumbago Proboscidea Stylidium See also International Carnivorous Plant Society Insectivorous Plant Society North American Sarracenia Conservancy List of carnivorous plants List of carnivorous plant periodicals Pitcher plant

Taxon identifiers Pinguicula Wikidata: Q161663 Wikispecies: Pinguicula APDB: 193740 BioLib: 40965 CoL: 6QMW EoL: 60117 EPPO: 1PNUG FNA: 125513 FoC: 125513 GBIF: 3172450 GRIN: 9414 iNaturalist: 72300 IPNI: 23916-1 IRMNG: 1084598 ITIS: 34433 NBN: NHMSYS0000461696 NCBI: 4197 NZOR: d249c856-b82c-4a1e-aa24-4700451f6fd6 Open Tree of Life: 659715 PLANTS: PINGU POWO: urn:lsid:ipni.org:names:30014019-2 Tropicos: 40009106 VASCAN: 1552 WFO: wfo-4000029780

Authority control databases International GND National United States Czech Republic Israel Other Yale LUX

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Adapted from the Wikipedia article [Pinguicula](https://en.wikipedia.org/wiki/Pinguicula) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Pinguicula?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.