Pinguicula moranensis
Encyclopedia
Pinguicula moranensis is a perennial rosette
-forming insectivorous
herb
native to Mexico
and Guatemala
. A species of butterwort, it forms summer rosettes
of flat, succulent leaves up to 10 centimeters (4 in) long, which are covered in mucilage
nous (sticky) glands that attract, trap, and digest arthropod
prey. Nutrients derived from the prey are used to supplement the nutrient-poor substrate
that the plant grows in. In the winter the plant forms a non-carnivorous rosette of small, fleshy leaves that conserves energy while food and moisture supplies are low. Single pink, purple, or violet flowers appear twice a year on upright stalks up to 25 centimeters long.
The species was first collected by Humboldt
and Bonpland
on the outskirts of Mina de Morán in the Sierra de Pachuca of the modern-day Mexican state of Hidalgo on their Latin American expedition of 1799–1804. Based on these collections, Humboldt, Bonpland and Carl Sigismund Kunth
described this species in Nova Genera et Species Plantarum in 1817. The extremely variable species has been redefined at least twice since, while several new species have been segregated from it based on various geographical or morphological
distinctions, although the legitimacy of some of these is still debated. P. moranensis remains the most common and most widely distributed member of the Section Orcheosanthus. It has long been cultivated for its carnivorous nature and attractive flowers, and is one of the most common butterworts in cultivation.
The generic name Pinguicula is derived from the Latin pinguis (meaning "fat") due to the buttery texture of the surface of the carnivorous leaves. The specific epithet moranensis refers to its type location, Mina de Moran.
leaves, each up to 95 millimeters (3¾ in) long. These leaves are carnivorous, having a large surface area densely covered with stalked mucilagenous glands with which they attract, trap, and digest arthropod prey, most commonly flies
. These so-called "summer leaves" are replaced by "winter rosettes" of small, glandless succulent leaves with the onset of the dry season in October. This protective winter rosette allows the plant to undergo winter dormancy
until the first rains begin in May. Flowers born singly on upright 10–25 centimeters (4–10 in.) peduncle
s emerge twice during the year (from the summer rosette and again from the winter rosette), a feature rare among the Mexican species. In the summer these appear in June, peak in August and September, and disappear with the return to the winter rosette in October or November.
The leaf blades of the summer rosettes of P. moranensis are smooth, rigid, and succulent, varying from bright yellow-green to maroon in colour. The laminae
are generally obovate
to orbicular
, between 5.5 and 13 centimeters (2–5 in.) long and supported by a 1 to 3.5 centimetre (⅜–1 ⅜ in.) petiole
.
As with all members of the genus, these leaf blades are densely covered by peduncular (stalked) mucilage
nous gland
s and sessile
(flat) digestive glands. The peduncular glands consist of a few secretory cells on top of a single-celled stalk. These cells produce a mucilagenous 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 sundew
s. The droplets secrete only limited 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. The insect struggles, triggering more glands and encasing itself in mucilage. P. moranensis can bend its leaf edges slightly by thigmotropism
, bringing additional glands into contact with the trapped insect. The sessile glands, which lie flat on the leaf surface, serve to digest the insect prey. Once the prey is entrapped by the peduncular glands and digestion begins, the initial flow of nitrogen triggers enzyme release by the sessile glands. These enzymes, which include amylase
, esterase
, phosphatase
, protease
, and ribonuclease
break down the digestible components of the insect body. These fluids are then absorbed back into the leaf surface through cuticular
holes, leaving only the chitin
exoskeleton
of the larger insects on the leaf surface.
The holes in the cuticle
which allow for this digestive mechanism pose a challenge for the plant, since they serve as breaks in the cuticle (waxy layer) that protects the plant from desiccation. As a result, P. moranensis is usually found in relatively humid environments. The production of the stalked capture glands and sessile digestive glands is also costly. A recent study found that the density of these respective glands can be correlated to environmental gradients. For example, capture gland density was found to be highest where prey availability was low, whereas digestive glands density showed direct correlation to prey availability. These results suggest that the amount of investment in carnivorous features is an adaptation to the environmental gradients.
" rosette of P. moranensis is two to three (maximum five) centimeters (¾–2 in.) in diameter and consists of 60 to 100 or more small, fleshy, non-glandular leaves. These are each 10 to 30 millimeters (⅜–1 ¼ in.) long and three to eight millimeters (⅛– in.) wide, generally spatulate
or oblong-spatulate, and densely covered with fine hairs
. The rosette is either open or compact and bulb-like, depending on variety
(see below).
s during each flowering period. These are borne singly on upright flower stalks which are green to brown-green in color and usually, like the upper surface of the carnivorous leaves, are densely covered in glandular hairs; the peduncles do, in fact, trap insect prey. The peduncles are 10 to 25 centimeters (4–10 in.) long and taper from two to three millimeters (⅛ in.) at the base to one millimeter ( in.) at the top.
The flowers themselves are composed of five petal
s which are fused at one end. The throat, the portion of the flower near the attachment point which holds the reproductive organs, is funnel shaped, and the petals flare out from there into a five-lobed zygomorphic corolla. The flowers 30 to 50 millimeters (1 ¼–2 in.) long. Below the attachment point to the stem the petals are fused into a 15–30 millimeter long spur which protrudes backwards roughly perpendicular to the rest of the flower.
The ovary and attached pistil protrude from the top of the floral tube near its opening, with the receptive stigma surface toward the front. Two 1 millimeter anthers hang from recurved, 2 millimeter filaments behind the pistil. Pollinators exiting after collecting nectar from the spur brush against the anther, transferring pollen to the stigma of the next flower they visit. The flowers can last up to 10 days but will wilt once they are pollinated. Pollinated ovaries ripen into 5 millimeter ( in.) dehiscent seed capsules containing numerous 1 millimeter long seeds. The chromosome count for this species is 2n=44.
The color and morphology of the flowers of this species is extremely variable, a source of delight to the horticulturist
and headache to the taxonomist
. Some generalizations, however, can be made.
The corolla flares open into five lobes, two upper lobes and three lower lobes. The upper lobes are 7–16 millimeters (¼–⅜ in.) long by 4–9 millimeters (–⅜ in.) wide and generally oblong, obovate, or cuneate. The lower lobes are similarly shaped and are 7–20 millimeters (¼–¾ in.) long by 4–18 millimeters (–¾ in.) wide. The central lower lobe is usually slightly longer than its neighbors. All of the petal lobes have rounded ends. The floral tube that houses the reproductive organs and is visible at the base of corolla lobes is white or lilac in color and 4–6 millimeters (–¼ in.) long. The white color of the floral tube can extend to a variable distance onto the corolla lobes, particularly in a stripe on the lower middle lobe. The color of the corolla lobes generally varies from pink to purple, but has been described by collectors as being "purple, scarlet, rosy-lavender to bluish-purple, dark pink to lavender, pinkish-purple, deep violet-purple, dark purple, bright mauve-pink, bright purple-pink, magenta with [white eye], [and] reddish pale with white eye." A rare white-flowered form is also known.
and Aimé Bonpland
's Latin American expedition in 1799–1804, only 8 Pinguicula species were known to science — 5 from Europe, 2 from North America and P. involuta from Peru
. From 1803–1805, three additional species from Europe and North America were described, bringing the total of known species to 11. In 1817 Humboldt
, Bonpland
and Carl Sigismund Kunth
described 3 new species from their Latin American expedition: The Peruvian P. calyptrata and the first known Mexican species: P. macrophylla and P. moranensis. At this point no infrageneric classification had yet been suggested.
In 1844 a French-Swiss botanist by the name of Alphonse Pyrame de Candolle (who created the first Code of Botanical Nomenclature) proposed a division of the genus into three sections
based on floral morphology
. He placed in the section Orcheosanthus those species with purple, deeply bilabiate corollas with 5 sub-equal lobes, a short floral tube, and a large spur not protruding past this tube. He included four species in this section, all of them from Mexico: P. oblongiloba, P. orchidioides
, P. caudata and P. moranensis. He excluded P. macrophylla H.B.K. on the grounds that it was a "dubious species".
The section Orcheosanthus grew as Charles Morren
described P. flos-mulionis in 1872, Eugene Fournier added P. sodalium in 1873 and Sander proposed P. bakeriana in 1881. In 1879–1888, however, a botanist by the name of William Hemsley
, after studying multiple specimens in herbariums and in culture, came to the conclusion that all the taxa placed in the section Orcheosanthus up to that point belonged to the same single species. Due to doubts as to the identity of the original two species described by H.
B.
K.
, Hemsley decided to use the name P. caudata Schltdl. for his conglomerate species. This name has been "indiscriminately" applied to members of the complex ever since.
in 1916, he recognized six species in the section Orcheosanthus, admitting however that this number was likely to change as others studied the section in the future. Sprague in 1928 proposed that the species joined by Hemsley were probably distinct, but that they were likely so interrelated that distinguishing between them would require observing characteristics that were usually or always indistinct in dried specimens. Sprague recognized eight species in the section: P. moranensis H.B.K, P. caudata Schltdl., P. oblongiloba, P. flos-mulionis, P. bakeriana, a P. moranensis-like P. rosei described by Watson in 1911, and the very distinct P. gypsicola
.
In 1966 Casper
published the first ever monograph of the genus. He clearly defined his taxonomic organization according to a broad range of morphological and phenotypic characteristics. Casper considered P. caudata, as well as various other taxa, to be synonyms of P. moranensis. He therefore recognized only 6 species for the section Orcheosanthus: P. moranensis, P. gypsicola, P. macrophylla H.B.K., P. oblongiloba, and the two recently discovered species P. colimensis and P. cyclosecta. Since that time 14 additional species have been discovered and assigned to the section. When Zamudio redefined the section in 1999, however, he chose to include only 12 species, including all six of Casper's choices. P. moranensis, therefore, remains in the section Orcheosanthus, along with over a dozen synonyms it has inherited in its 200-year taxonomic history.
Pinguicula. Using molecular data, they were able to isolate those morphological characteristics that were synapomorphies
for various groups, providing evidence for a genetically based taxonomic structure. Their overall results did not support the placement of P. moranensis in the section Orcheosanthus, rather indicating that it should be placed in the section Longitubus along with P. laueana.
In further disagreement with Ruiz's 2001 revision of the section Orcheosanthus, Cieslak et al.'s phylogenetic data indicated that P. rectifolia and several unnamed taxa that had been treated as synonyms of P. moranensis are in fact a distinct complex. They isolated several morphological characteristics that could be used to differentiate between the complexes, including floral spur length (longer in P. moranensis), flower color (never with a blue tinge in P. moranensis), and the shape of the lateral corolla lobe (exhibiting a twist in P. rectifolia). A more thorough study analyzing numerous P. moranensis populations and other members of closely related taxa is needed to resolve this complex.
, mainly on the basis of the shape of the leaves composing their winter (resting) rosettes:
This variety has open winter rosettes composed of leaves which are spatulate in shape and have an obtuse or rounded end. It tends to grow in limestone-based substrates.
This variety has a closed, bulb-like rosette of winter leaves which are acicular (pointy) at the tip. It tends to grow on igneous substrates.
Ruiz also noted that these subspecies differed in their preference of soil substrate. He first noted this while attempting to find the population of plants from which Humboldt
and Bonpland
had collected their type specimens in 1803. Although Ruiz was able to find many populations of the species growing in the areas that Humboldt and Bonpland had visited in the vicinity if Mina de Moran, only one population, the only one growing on limestone, matched H.B.K.'s description and their type and isotype specimens now housed in the herbarium
of the French national museum of natural history
. The other populations in the area grew on substrate of igneous origin and more closely matched Hooker's
1846 description of P. orchidioides. These latter plants then became the new variety, P. moranensis ssp. neovolcanica.
P. moranensis is the most widely distributed member of the Section Orcheosanthus. It is also the most common and widely distributed Pinguicula species in Mexico, being found in all the major mountain ranges except Sierra Madre Occidental and Baja California. Locations are known from the Mexican states of Tamaulipas
, Guanajuato
, Nuevo León
, Campeche
, Chiapas
, Oaxaca
, Puebla
, Distrito Federal, Veracruz
, Estado de México, Querétaro
, San Luis Potosí
, Morelos
, Hidalgo, Guerrero
, Zacatecas
, Tlaxcala
, Quintana Roo
, and Michoacán
and the Guatemala
n departments
of Huehuetenango
, Quiché
, San Marcos
, Quetzaltenango
, Totonicapan
, Solola
, Chimaltenango
, Baja Verapaz
, Guatemala
and El Progreso
. Here it grows in mountainous regions between 800 and 3200 meters (2600–10500 ft) in altitude. Generally, the species tends to follow sedimentary outcrops of the Cretaceous
period. P. moranensis var. neovolcanica, however, tends to grow on igneous rocks of the Eje Volcánico Transversal.
P. moranensis most often grows in oak, pine-oak
, or temperate montane woodlands. However, its distribution penetrates into tropical forests and xerophytic
shrublands, as well as in gorges and canyon walls with high environmental humidity
. P. moranensis prefers humid and shady environments, such as slopes by streams, gullies, or road cuts, or among leaf litter in sandy soil high in organic matter. Its ability to gather nutrients from the arthropod prey it catches allows it to grow in low-nutrient environments where other plants would usually out-compete it. As a result, it is often found in disturbed areas or on steep cliffs or hillsides. Since its roots do little more than provide anchorage, the plant requires little or no soil, and dense clusters can be found clinging onto boulders, moss or crags in rock faces, or even epiphytically
on tree trunks. Common companion plants include moss
es, Selaginella, fern
s and other herbaceous plants, as well as canopy trees such as pines and oaks.
, perlite
, vermiculite
, peat moss, gypsum
and/or decomposed granite. Soil should be kept well drained, but watered regularly with distilled water
in the summer and only very rarely once the plant enters its winter rosette. The species grows readily on well-lit windowsills, under fluorescent lights, or in warm to hot greenhouses.
s involving the species have been registered and are recognized by the International Carnivorous Plant Society
:
Additionally, three clones of P. moranensis have been registered as cultivars:
Rosette (botany)
In botany, a rosette is a circular arrangement of leaves, with all the leaves at a single height.Though rosettes usually sit near the soil, their structure is an example of a modified stem.-Function:...
-forming insectivorous
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. Carnivorous plants appear adapted to grow in places where the soil is thin or poor in nutrients, especially nitrogen, such as acidic...
herb
Herbaceous
A herbaceous plant is a plant that has leaves and stems that die down at the end of the growing season to the soil level. They have no persistent woody stem above ground...
native to Mexico
Mexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...
and Guatemala
Guatemala
Guatemala is a country in Central America bordered by Mexico to the north and west, the Pacific Ocean to the southwest, Belize to the northeast, the Caribbean to the east, and Honduras and El Salvador to the southeast...
. A species of butterwort, it forms summer rosettes
Rosette (botany)
In botany, a rosette is a circular arrangement of leaves, with all the leaves at a single height.Though rosettes usually sit near the soil, their structure is an example of a modified stem.-Function:...
of flat, succulent leaves up to 10 centimeters (4 in) long, which are covered in mucilage
Mucilage
Mucilage is a thick, gluey substance produced by most plants and some microorganisms. It is a polar glycoprotein and an exopolysaccharide.It occurs in various parts of nearly all classes of plant, usually in relatively small percentages, and is frequently associated with other substances, such as...
nous (sticky) glands that attract, trap, and digest arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...
prey. Nutrients derived from the prey are used to supplement the nutrient-poor substrate
Substrate (biology)
In biology a substrate is the surface a plant or animal lives upon and grows on. A substrate can include biotic or abiotic materials and animals. For example, encrusting algae that lives on a rock can be substrate for another animal that lives on top of the algae. See also substrate .-External...
that the plant grows in. In the winter the plant forms a non-carnivorous rosette of small, fleshy leaves that conserves energy while food and moisture supplies are low. Single pink, purple, or violet flowers appear twice a year on upright stalks up to 25 centimeters long.
The species was first collected by Humboldt
Alexander von Humboldt
Friedrich Wilhelm Heinrich Alexander Freiherr von Humboldt was a German naturalist and explorer, and the younger brother of the Prussian minister, philosopher and linguist Wilhelm von Humboldt...
and Bonpland
Aimé Bonpland
Aimé Jacques Alexandre Bonpland was a French explorer and botanist.Bonpland's real name was Goujaud, and he was born in La Rochelle, a coastal city in France. After serving as a surgeon in the French army, and studying under J. N...
on the outskirts of Mina de Morán in the Sierra de Pachuca of the modern-day Mexican state of Hidalgo on their Latin American expedition of 1799–1804. Based on these collections, Humboldt, Bonpland and Carl Sigismund Kunth
Carl Sigismund Kunth
Carl Sigismund Kunth , also Karl Sigismund Kunth or anglicized as Charles Sigismund Kunth, was a German botanist...
described this species in Nova Genera et Species Plantarum in 1817. The extremely variable species has been redefined at least twice since, while several new species have been segregated from it based on various geographical or morphological
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
distinctions, although the legitimacy of some of these is still debated. P. moranensis remains the most common and most widely distributed member of the Section Orcheosanthus. It has long been cultivated for its carnivorous nature and attractive flowers, and is one of the most common butterworts in cultivation.
The generic name Pinguicula is derived from the Latin pinguis (meaning "fat") due to the buttery texture of the surface of the carnivorous leaves. The specific epithet moranensis refers to its type location, Mina de Moran.
Plant characteristics
Habit
P. moranensis is seasonally dimorphic, in that it undergoes two distinct growth habits throughout the year. During the summer when rain and insect prey are most plentiful, the plant forms a ground hugging rosette composed of 6–8 generally obovateLeaf shape
In botany, leaf shape is characterised with the following terms :* Acicular : Slender and pointed, needle-like* Acuminate : Tapering to a long point...
leaves, each up to 95 millimeters (3¾ in) long. These leaves are carnivorous, having a large surface area densely covered with stalked mucilagenous glands with which they attract, trap, and digest arthropod prey, most commonly flies
Diptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
. These so-called "summer leaves" are replaced by "winter rosettes" of small, glandless succulent leaves with the onset of the dry season in October. This protective winter rosette allows the plant to undergo winter dormancy
Dormancy
Dormancy is a period in an organism's life cycle when growth, development, and physical activity are temporarily stopped. This minimizes metabolic activity and therefore helps an organism to conserve energy. Dormancy tends to be closely associated with environmental conditions...
until the first rains begin in May. Flowers born singly on upright 10–25 centimeters (4–10 in.) peduncle
Peduncle (botany)
In botany, a peduncle is a stem supporting an inflorescence, or after fecundation, an infructescence.The peduncle is a stem, usually green and without leaves, though sometimes colored or supporting small leaves...
s emerge twice during the year (from the summer rosette and again from the winter rosette), a feature rare among the Mexican species. In the summer these appear in June, peak in August and September, and disappear with the return to the winter rosette in October or November.
Leaves and carnivory
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....
are generally obovate
Leaf shape
In botany, leaf shape is characterised with the following terms :* Acicular : Slender and pointed, needle-like* Acuminate : Tapering to a long point...
to orbicular
Leaf shape
In botany, leaf shape is characterised with the following terms :* Acicular : Slender and pointed, needle-like* Acuminate : Tapering to a long point...
, between 5.5 and 13 centimeters (2–5 in.) long and supported by a 1 to 3.5 centimetre (⅜–1 ⅜ in.) petiole
Petiole (botany)
In botany, the petiole is the stalk attaching the leaf blade to the stem. The petiole usually has the same internal structure as the stem. Outgrowths appearing on each side of the petiole are called stipules. Leaves lacking a petiole are called sessile, or clasping when they partly surround the...
.
As with all members of the genus, these leaf blades are densely covered by peduncular (stalked) mucilage
Mucilage
Mucilage is a thick, gluey substance produced by most plants and some microorganisms. It is a polar glycoprotein and an exopolysaccharide.It occurs in various parts of nearly all classes of plant, usually in relatively small percentages, and is frequently associated with other substances, such as...
nous gland
Gland
A gland is an organ in an animal's body that synthesizes a substance for release of substances such as hormones or breast milk, often into the bloodstream or into cavities inside the body or its outer surface .- Types :...
s and sessile
Sessility (botany)
In botany, sessility is a characteristic of plants whose flowers or leaves are borne directly from the stem or peduncle, and thus lack a petiole or pedicel...
(flat) digestive glands. The peduncular glands consist of a few secretory cells on top of a single-celled stalk. These cells produce a mucilagenous 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 sundew
Sundew
Drosera, commonly known as the sundews, comprise one of the largest genera of carnivorous plants, with at least 194 species. These members of the family Droseraceae lure, capture, and digest insects using stalked mucilaginous glands covering their leaf surface. The insects are used to supplement...
s. The droplets secrete only limited 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. The insect struggles, triggering more glands and encasing itself in mucilage. P. moranensis can bend its leaf edges slightly by thigmotropism
Thigmotropism
Thigmotropism is a movement in which an organism moves or grows in response to touch or contact stimuli. The prefix thigmo- θιγμος comes from the Greek for "touch". Usually thigmotropism occurs when plants grow around a surface, such as a wall, pot, or trellis. Climbing plants, such as vines,...
, bringing additional glands into contact with the trapped insect. The sessile glands, which lie flat on the leaf surface, serve to digest the insect prey. Once the prey is entrapped by the peduncular glands and digestion begins, the initial flow of nitrogen triggers enzyme release by the sessile glands. These enzymes, which include amylase
Amylase
Amylase is an enzyme that catalyses the breakdown of starch into sugars. Amylase is present in human saliva, where it begins the chemical process of digestion. Food that contains much starch but little sugar, such as rice and potato, taste slightly sweet as they are chewed because amylase turns...
, esterase
Esterase
An esterase is a hydrolase enzyme that splits esters into an acid and an alcohol in a chemical reaction with water called hydrolysis.A wide range of different esterases exist that differ in their substrate specificity, their protein structure, and their biological function.- EC classification/list...
, phosphatase
Phosphatase
A phosphatase is an enzyme that removes a phosphate group from its substrate by hydrolysing phosphoric acid monoesters into a phosphate ion and a molecule with a free hydroxyl group . This action is directly opposite to that of phosphorylases and kinases, which attach phosphate groups to their...
, protease
Protease
A protease is any enzyme that conducts proteolysis, that is, begins protein catabolism by hydrolysis of the peptide bonds that link amino acids together in the polypeptide chain forming the protein....
, and ribonuclease
Ribonuclease
Ribonuclease 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 EC 2.7 and 3.1 classes of enzymes.-Function:All organisms studied contain...
break down the digestible components of the insect body. These fluids are then absorbed back into the leaf surface through cuticular
Plant cuticle
Plant cuticles are a protective waxy covering produced only by the epidermal cells of leaves, young shoots and all other aerial plant organs without periderm...
holes, leaving only the chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
of the larger insects on the leaf surface.
The holes in the cuticle
Plant cuticle
Plant cuticles are a protective waxy covering produced only by the epidermal cells of leaves, young shoots and all other aerial plant organs without periderm...
which allow for this digestive mechanism pose a challenge for the plant, since they serve as breaks in the cuticle (waxy layer) that protects the plant from desiccation. As a result, P. moranensis is usually found in relatively humid environments. The production of the stalked capture glands and sessile digestive glands is also costly. A recent study found that the density of these respective glands can be correlated to environmental gradients. For example, capture gland density was found to be highest where prey availability was low, whereas digestive glands density showed direct correlation to prey availability. These results suggest that the amount of investment in carnivorous features is an adaptation to the environmental gradients.
Winter rosette
The "winter" or "restingEstivation
Aestivation is a state of animal dormancy, characterized by inactivity and a lowered metabolic rate, that is entered in response to high temperatures and arid conditions...
" rosette of P. moranensis is two to three (maximum five) centimeters (¾–2 in.) in diameter and consists of 60 to 100 or more small, fleshy, non-glandular leaves. These are each 10 to 30 millimeters (⅜–1 ¼ in.) long and three to eight millimeters (⅛– in.) wide, generally spatulate
Leaf shape
In botany, leaf shape is characterised with the following terms :* Acicular : Slender and pointed, needle-like* Acuminate : Tapering to a long point...
or oblong-spatulate, and densely covered with fine hairs
Trichome
Trichomes are fine outgrowths or appendages on plants and certain protists. These are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae.- Algal trichomes :...
. The rosette is either open or compact and bulb-like, depending on variety
Variety (biology)
In botanical nomenclature, variety is a taxonomic rank below that of species: as such, it gets a three-part infraspecific name....
(see below).
Flowers
P. moranensis produces one to seven flowerFlower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...
s during each flowering period. These are borne singly on upright flower stalks which are green to brown-green in color and usually, like the upper surface of the carnivorous leaves, are densely covered in glandular hairs; the peduncles do, in fact, trap insect prey. The peduncles are 10 to 25 centimeters (4–10 in.) long and taper from two to three millimeters (⅛ in.) at the base to one millimeter ( in.) at the top.
Petal
Petals are modified leaves that surround the reproductive parts of flowers. They often are brightly colored or unusually shaped to attract pollinators. Together, all of the petals of a flower are called a corolla. Petals are usually accompanied by another set of special leaves called sepals lying...
s which are fused at one end. The throat, the portion of the flower near the attachment point which holds the reproductive organs, is funnel shaped, and the petals flare out from there into a five-lobed zygomorphic corolla. The flowers 30 to 50 millimeters (1 ¼–2 in.) long. Below the attachment point to the stem the petals are fused into a 15–30 millimeter long spur which protrudes backwards roughly perpendicular to the rest of the flower.
The ovary and attached pistil protrude from the top of the floral tube near its opening, with the receptive stigma surface toward the front. Two 1 millimeter anthers hang from recurved, 2 millimeter filaments behind the pistil. Pollinators exiting after collecting nectar from the spur brush against the anther, transferring pollen to the stigma of the next flower they visit. The flowers can last up to 10 days but will wilt once they are pollinated. Pollinated ovaries ripen into 5 millimeter ( in.) dehiscent seed capsules containing numerous 1 millimeter long seeds. The chromosome count for this species is 2n=44.
Horticulture
Horticulture is the industry and science of plant cultivation including the process of preparing soil for the planting of seeds, tubers, or cuttings. Horticulturists work and conduct research in the disciplines of plant propagation and cultivation, crop production, plant breeding and genetic...
and headache to the taxonomist
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...
. Some generalizations, however, can be made.
The corolla flares open into five lobes, two upper lobes and three lower lobes. The upper lobes are 7–16 millimeters (¼–⅜ in.) long by 4–9 millimeters (–⅜ in.) wide and generally oblong, obovate, or cuneate. The lower lobes are similarly shaped and are 7–20 millimeters (¼–¾ in.) long by 4–18 millimeters (–¾ in.) wide. The central lower lobe is usually slightly longer than its neighbors. All of the petal lobes have rounded ends. The floral tube that houses the reproductive organs and is visible at the base of corolla lobes is white or lilac in color and 4–6 millimeters (–¼ in.) long. The white color of the floral tube can extend to a variable distance onto the corolla lobes, particularly in a stripe on the lower middle lobe. The color of the corolla lobes generally varies from pink to purple, but has been described by collectors as being "purple, scarlet, rosy-lavender to bluish-purple, dark pink to lavender, pinkish-purple, deep violet-purple, dark purple, bright mauve-pink, bright purple-pink, magenta with [white eye], [and] reddish pale with white eye." A rare white-flowered form is also known.
Taxonomy
Sergio Zamudio Ruiz, in his 2001 revision of the section Orcheosanthus, called the identity and exact delimitation of P. moranensis "perhaps the most difficult problem to solve within the genus". This difficulty is due mainly to the high variability and large geographic distribution of the species, which has given rise to the description of many synonyms since the species was first described nearly 200 years ago. Botanists have attempted to delimitate the species through various morphological, ecological and genetic methods, though to this date some debate remains as to the placement and description of P. moranensis and its relationship to the species to which it is closely related.Botanical history
Prior to Alexander von HumboldtAlexander von Humboldt
Friedrich Wilhelm Heinrich Alexander Freiherr von Humboldt was a German naturalist and explorer, and the younger brother of the Prussian minister, philosopher and linguist Wilhelm von Humboldt...
and Aimé Bonpland
Aimé Bonpland
Aimé Jacques Alexandre Bonpland was a French explorer and botanist.Bonpland's real name was Goujaud, and he was born in La Rochelle, a coastal city in France. After serving as a surgeon in the French army, and studying under J. N...
's Latin American expedition in 1799–1804, only 8 Pinguicula species were known to science — 5 from Europe, 2 from North America and P. involuta from Peru
Peru
Peru , officially the Republic of Peru , is a country in western South America. It is bordered on the north by Ecuador and Colombia, on the east by Brazil, on the southeast by Bolivia, on the south by Chile, and on the west by the Pacific Ocean....
. From 1803–1805, three additional species from Europe and North America were described, bringing the total of known species to 11. In 1817 Humboldt
Alexander von Humboldt
Friedrich Wilhelm Heinrich Alexander Freiherr von Humboldt was a German naturalist and explorer, and the younger brother of the Prussian minister, philosopher and linguist Wilhelm von Humboldt...
, Bonpland
Aimé Bonpland
Aimé Jacques Alexandre Bonpland was a French explorer and botanist.Bonpland's real name was Goujaud, and he was born in La Rochelle, a coastal city in France. After serving as a surgeon in the French army, and studying under J. N...
and Carl Sigismund Kunth
Carl Sigismund Kunth
Carl Sigismund Kunth , also Karl Sigismund Kunth or anglicized as Charles Sigismund Kunth, was a German botanist...
described 3 new species from their Latin American expedition: The Peruvian P. calyptrata and the first known Mexican species: P. macrophylla and P. moranensis. At this point no infrageneric classification had yet been suggested.
In 1844 a French-Swiss botanist by the name of Alphonse Pyrame de Candolle (who created the first Code of Botanical Nomenclature) proposed a division of the genus into three sections
Section (botany)
In botany, a section is a taxonomic rank below the genus, but above the species. The subgenus, if present, is higher than the section, and the rank of series, if present, is below the section. Sections are typically used to help organise very large genera, which may have hundreds of species...
based on floral morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
. He placed in the section Orcheosanthus those species with purple, deeply bilabiate corollas with 5 sub-equal lobes, a short floral tube, and a large spur not protruding past this tube. He included four species in this section, all of them from Mexico: P. oblongiloba, P. orchidioides
Pinguicula orchidioides
Pinguicula orchidioides is a perennial rosette-forming insectivorous herb native to Mexico and Guatemala. A species of butterwort, it forms summer rosettes of flat, succulent leaves up to 5 centimeters long, which are covered in mucilagenous glands that attract, trap, and digest arthropod...
, P. caudata and P. moranensis. He excluded P. macrophylla H.B.K. on the grounds that it was a "dubious species".
Charles Jacques Édouard Morren
Charles Jacques Édouard Morren , was a Belgian botanist, professor of botany and director of the Jardin botanique de l'Université de Liège from 1857-1886. His special field of study was the Bromeliaceae on which family he was the recognised authority...
described P. flos-mulionis in 1872, Eugene Fournier added P. sodalium in 1873 and Sander proposed P. bakeriana in 1881. In 1879–1888, however, a botanist by the name of William Hemsley
William Hemsley (botanist)
William Botting Hemsley was an English botanist and 1909 Victoria Medal of Honour recipient....
, after studying multiple specimens in herbariums and in culture, came to the conclusion that all the taxa placed in the section Orcheosanthus up to that point belonged to the same single species. Due to doubts as to the identity of the original two species described by H.
Alexander von Humboldt
Friedrich Wilhelm Heinrich Alexander Freiherr von Humboldt was a German naturalist and explorer, and the younger brother of the Prussian minister, philosopher and linguist Wilhelm von Humboldt...
B.
Aimé Bonpland
Aimé Jacques Alexandre Bonpland was a French explorer and botanist.Bonpland's real name was Goujaud, and he was born in La Rochelle, a coastal city in France. After serving as a surgeon in the French army, and studying under J. N...
K.
Carl Sigismund Kunth
Carl Sigismund Kunth , also Karl Sigismund Kunth or anglicized as Charles Sigismund Kunth, was a German botanist...
, Hemsley decided to use the name P. caudata Schltdl. for his conglomerate species. This name has been "indiscriminately" applied to members of the complex ever since.
Twentieth century
When Barnhart revised the family LentibulariaceaeLentibulariaceae
Lentibulariaceae is a family of carnivorous plants containing three genera, Genlisea, the corkscrew plants, Pinguicula, the butterworts, and Utricularia, the bladderworts....
in 1916, he recognized six species in the section Orcheosanthus, admitting however that this number was likely to change as others studied the section in the future. Sprague in 1928 proposed that the species joined by Hemsley were probably distinct, but that they were likely so interrelated that distinguishing between them would require observing characteristics that were usually or always indistinct in dried specimens. Sprague recognized eight species in the section: P. moranensis H.B.K, P. caudata Schltdl., P. oblongiloba, P. flos-mulionis, P. bakeriana, a P. moranensis-like P. rosei described by Watson in 1911, and the very distinct P. gypsicola
Pinguicula gypsicola
Pinguicula gypsicola is an insectivorous plant of the genus Pinguicula native to the Mexican state of San Luis Potosi, a heterophyllous member of the section Orcheosanthus. It grows in gypsum soils and forms stemless rosettes of upright, narrow leaves.-Morphology:P. gypsicola is a perennial...
.
In 1966 Casper
Siegfried Jost Casper
Siegfried Jost Casper is a German biologist whose primary research is in limnology and the plant genus Pinguicula . Together with Heinz-Dieter Krausch he has published a basic reference work on the sweet-water flora of central Europe. For many years he studied the East German lake Stechlinsee as...
published the first ever monograph of the genus. He clearly defined his taxonomic organization according to a broad range of morphological and phenotypic characteristics. Casper considered P. caudata, as well as various other taxa, to be synonyms of P. moranensis. He therefore recognized only 6 species for the section Orcheosanthus: P. moranensis, P. gypsicola, P. macrophylla H.B.K., P. oblongiloba, and the two recently discovered species P. colimensis and P. cyclosecta. Since that time 14 additional species have been discovered and assigned to the section. When Zamudio redefined the section in 1999, however, he chose to include only 12 species, including all six of Casper's choices. P. moranensis, therefore, remains in the section Orcheosanthus, along with over a dozen synonyms it has inherited in its 200-year taxonomic history.
Phylogenetics
The varying importance which different authors have placed on various morphological characteristics when determining the taxonomy of the genus has long made the resulting subdivision of the genus a subject of controversy. Ruiz (2001) supported his revision of the section Orcheosanthus with a phylogenetic analysis, using 20 morphological and phenological characteristics. In 2005, Cieslak et al. conducted the first comprehensive phylogenetic analysis of the entire genusGenus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
Pinguicula. Using molecular data, they were able to isolate those morphological characteristics that were synapomorphies
Synapomorphy
In cladistics, a synapomorphy or synapomorphic character is a trait that is shared by two or more taxa and their most recent common ancestor, whose ancestor in turn does not possess the trait. A synapomorphy is thus an apomorphy visible in multiple taxa, where the trait in question originates in...
for various groups, providing evidence for a genetically based taxonomic structure. Their overall results did not support the placement of P. moranensis in the section Orcheosanthus, rather indicating that it should be placed in the section Longitubus along with P. laueana.
In further disagreement with Ruiz's 2001 revision of the section Orcheosanthus, Cieslak et al.'s phylogenetic data indicated that P. rectifolia and several unnamed taxa that had been treated as synonyms of P. moranensis are in fact a distinct complex. They isolated several morphological characteristics that could be used to differentiate between the complexes, including floral spur length (longer in P. moranensis), flower color (never with a blue tinge in P. moranensis), and the shape of the lateral corolla lobe (exhibiting a twist in P. rectifolia). A more thorough study analyzing numerous P. moranensis populations and other members of closely related taxa is needed to resolve this complex.
Varieties
After extensively studying P. moranensis in habitat, Ruiz (1999) came to the conclusion that the species could be divided into two distinct varietiesVariety (biology)
In botanical nomenclature, variety is a taxonomic rank below that of species: as such, it gets a three-part infraspecific name....
, mainly on the basis of the shape of the leaves composing their winter (resting) rosettes:
- Pinguicula moranensis Kunth in Humb.Alexander von HumboldtFriedrich Wilhelm Heinrich Alexander Freiherr von Humboldt was a German naturalist and explorer, and the younger brother of the Prussian minister, philosopher and linguist Wilhelm von Humboldt...
, Bonpl.Aimé BonplandAimé Jacques Alexandre Bonpland was a French explorer and botanist.Bonpland's real name was Goujaud, and he was born in La Rochelle, a coastal city in France. After serving as a surgeon in the French army, and studying under J. N...
et Kunth. var. moranensis
This variety has open winter rosettes composed of leaves which are spatulate in shape and have an obtuse or rounded end. It tends to grow in limestone-based substrates.
- Pinguicula moranensis Kunth in Humb.Alexander von HumboldtFriedrich Wilhelm Heinrich Alexander Freiherr von Humboldt was a German naturalist and explorer, and the younger brother of the Prussian minister, philosopher and linguist Wilhelm von Humboldt...
, Bonpl.Aimé BonplandAimé Jacques Alexandre Bonpland was a French explorer and botanist.Bonpland's real name was Goujaud, and he was born in La Rochelle, a coastal city in France. After serving as a surgeon in the French army, and studying under J. N...
et Kunth. var. neovolcanica
This variety has a closed, bulb-like rosette of winter leaves which are acicular (pointy) at the tip. It tends to grow on igneous substrates.
Ruiz also noted that these subspecies differed in their preference of soil substrate. He first noted this while attempting to find the population of plants from which Humboldt
Alexander von Humboldt
Friedrich Wilhelm Heinrich Alexander Freiherr von Humboldt was a German naturalist and explorer, and the younger brother of the Prussian minister, philosopher and linguist Wilhelm von Humboldt...
and Bonpland
Aimé Bonpland
Aimé Jacques Alexandre Bonpland was a French explorer and botanist.Bonpland's real name was Goujaud, and he was born in La Rochelle, a coastal city in France. After serving as a surgeon in the French army, and studying under J. N...
had collected their type specimens in 1803. Although Ruiz was able to find many populations of the species growing in the areas that Humboldt and Bonpland had visited in the vicinity if Mina de Moran, only one population, the only one growing on limestone, matched H.B.K.'s description and their type and isotype specimens now housed in the herbarium
Herbarium
In botany, a herbarium – sometimes known by the Anglicized term herbar – is a collection of preserved plant specimens. These specimens may be whole plants or plant parts: these will usually be in a dried form, mounted on a sheet, but depending upon the material may also be kept in...
of the French national museum of natural history
Muséum national d'histoire naturelle
The Muséum national d'Histoire naturelle is the National Museum of Natural History in Paris, France.- History :The museum was formally founded on 10 June 1793, during the French Revolution...
. The other populations in the area grew on substrate of igneous origin and more closely matched Hooker's
William Jackson Hooker
Sir William Jackson Hooker, FRS was an English systematic botanist and organiser. He held the post of Regius Professor of Botany at Glasgow University, and was the first Director of the Royal Botanic Gardens, Kew. He enjoyed the friendship and support of Sir Joseph Banks for his exploring,...
1846 description of P. orchidioides. These latter plants then became the new variety, P. moranensis ssp. neovolcanica.
Distribution and habitat
Tamaulipas
Tamaulipas officially Estado Libre y Soberano de Tamaulipas is one of the 31 states which, with the Federal District, comprise the 32 Federal Entities of Mexico. It is divided in 43 municipalities and its capital city is Ciudad Victoria. The capital city was named after Guadalupe Victoria, the...
, Guanajuato
Guanajuato
Guanajuato officially Estado Libre y Soberano de Guanajuato is one of the 31 states which, with the Federal District, comprise the 32 Federal Entities of Mexico. It is divided in 46 municipalities and its capital city is Guanajuato....
, Nuevo León
Nuevo León
Nuevo León It is located in Northeastern Mexico. It is bordered by the states of Tamaulipas to the north and east, San Luis Potosí to the south, and Coahuila to the west. To the north, Nuevo León has a 15 kilometer stretch of the U.S.-Mexico border adjacent to the U.S...
, Campeche
Campeche
Campeche is one of the 31 states which, with the Federal District, comprise the 32 Federal Entities of Mexico. Located in Southeast Mexico, it is bordered by the states of Yucatán to the north east, Quintana Roo to the east, and Tabasco to the south west...
, Chiapas
Chiapas
Chiapas officially Estado Libre y Soberano de Chiapas is one of the 31 states that, with the Federal District, comprise the 32 Federal Entities of Mexico. It is divided in 118 municipalities and its capital city is Tuxtla Gutierrez. Other important cites in Chiapas include San Cristóbal de las...
, Oaxaca
Oaxaca
Oaxaca , , officially Estado Libre y Soberano de Oaxaca is one of the 31 states which, along with the Federal District, comprise the 32 federative entities of Mexico. It is divided into 571 municipalities; of which 418 are governed by the system of customs and traditions...
, Puebla
Puebla
Puebla officially Estado Libre y Soberano de Puebla is one of the 31 states which, with the Federal District, comprise the 32 Federal Entities of Mexico. It is divided in 217 municipalities and its capital city is Puebla....
, Distrito Federal, Veracruz
Veracruz
Veracruz, formally Veracruz de Ignacio de la Llave officially Estado Libre y Soberano de Veracruz de Ignacio de la Llave , is one of the 31 states that, along with the Federal District, comprise the 32 federative entities of Mexico. It is divided in 212 municipalities and its capital city is...
, Estado de México, Querétaro
Querétaro
Querétaro officially Estado Libre y Soberano de Querétaro de Arteaga is one of the 31 states which, with the Federal District, comprise the 32 Federal Entities of Mexico. It is divided into 18 municipalities and its capital city is Santiago de Querétaro....
, San Luis Potosí
San Luis Potosí
San Luis Potosí officially Estado Libre y Soberano de San Luis Potosí is one of the 31 states which, with the Federal District, comprise the 32 Federal Entities of Mexico. It is divided in 58 municipalities and its capital city is San Luis Potosí....
, Morelos
Morelos
Morelos officially Estado Libre y Soberano de Morelos is one of the 31 states which, with the Federal District, comprise the 32 Federal Entities of Mexico. It is divided in 33 municipalities and its capital city is Cuernavaca....
, Hidalgo, Guerrero
Guerrero
Guerrero officially Estado Libre y Soberano de Guerrero is one of the 31 states which, with the Federal District, comprise the 32 Federal Entities of Mexico. It is divided in 81 municipalities and its capital city is Chilpancingo....
, Zacatecas
Zacatecas
Zacatecas officially Estado Libre y Soberano de Zacatecas is one of the 31 states which, with the Federal District, comprise the 32 Federal Entities of Mexico. It is divided in 58 municipalities and its capital city is Zacatecas....
, Tlaxcala
Tlaxcala
Tlaxcala officially Estado Libre y Soberano de Tlaxcala is one of the 31 states which along with the Federal District comprise the 32 federative entities of Mexico. It is divided into 60 municipalities and its capital city is Tlaxcala....
, Quintana Roo
Quintana Roo
Quintana Roo officially Estado Libre y Soberano de Quintana Roo is one of the 31 states which, with the Federal District, comprise the 32 Federal Entities of Mexico. It is divided in 10 municipalities and its capital city is Chetumal....
, and Michoacán
Michoacán
Michoacán officially Estado Libre y Soberano de Michoacán de Ocampo is one of the 31 states which, with the Federal District, comprise the 32 Federal Entities of Mexico. It is divided in 113 municipalities and its capital city is Morelia...
and the Guatemala
Guatemala
Guatemala is a country in Central America bordered by Mexico to the north and west, the Pacific Ocean to the southwest, Belize to the northeast, the Caribbean to the east, and Honduras and El Salvador to the southeast...
n departments
Departments of Guatemala
||Guatemala is divided into 22 departments :#Alta Verapaz#Baja Verapaz#Chimaltenango#Chiquimula#Petén#El Progreso#El Quiché#Escuintla#Guatemala#Huehuetenango#Izabal#Jalapa#Jutiapa#Quetzaltenango#Retalhuleu#Sacatepéquez...
of Huehuetenango
Huehuetenango
Huehuetenango is a city and a municipality in the highlands of western Guatemala. It is also the capital of the department of Huehuetenango. The municipality's population was over 81,000 people in 2002...
, Quiché
Quiché (department)
El Quiché is a department of Guatemala.El Quiché department is in the heartland of the Quiché people, to the north-west of Guatemala City. The capital is Santa Cruz del Quiché.-Population:...
, San Marcos
San Marcos, Guatemala
San Marcos San Marcos San Marcos (elevation: 7,868 feet (2,398 meters) is a city and municipality in Guatemala.It is the capital of the department of San Marcos.The municipality has a population of approximately 45,000.-Sports:...
, Quetzaltenango
Quetzaltenango
Quetzaltenango, also commonly known by its indigenous name, Xelajú , or more commonly, Xela , is the second largest city of Guatemala. It is both the capital of Quetzaltenango Department and the municipal seat of Quetzaltenango municipality....
, Totonicapan
Totonicapán
Totonicapán is a city in Guatemala. It serves as the capital of the department of Totonicapán and as the administrative seat for the surrounding municipality of Totonicapán.- External links :# #...
, Solola
Sololá
Sololá is a city in Guatemala. It is the capital of the department of Sololá and the administrative seat of Sololá municipality.The name is a hispanicized form of its pre-Columbian name, one spelling variant of which is T'zolojy'a...
, Chimaltenango
Chimaltenango
Chimaltenango is a town in Guatemala of some 43,900 people . It serves as both the capital of the department of Chimaltenango and the municipal seat for the surrounding municipality of the same name....
, Baja Verapaz
Baja Verapaz
Baja Verapaz is a department in Guatemala. The capital is Salamá.Baja Verapaz houses the Mario Dary Biotope Preserve, preserving the native flora and fauna of the region, especially the endangered national bird of Guatemala, the Resplendent Quetzal....
, Guatemala
Guatemala (department)
Guatemala is one of the 22 departments of Guatemala. The capital is Guatemala City, which also serves as the national capital.The department covers a surface area of 2,126 km², and had a population of 2,541,581 at the 2002 census.- Municipalities :...
and El Progreso
El Progreso (department)
El Progreso is a department in Guatemala. The capital is Guastatoya and the largest city is Sanarate.- Municipalities :# El Jícaro# Guastatoya# Morazán# San Agustín Acasaguastlán# San Antonio La Paz# San Cristóbal Acasaguastlán# Sanarate# Sansarate...
. Here it grows in mountainous regions between 800 and 3200 meters (2600–10500 ft) in altitude. Generally, the species tends to follow sedimentary outcrops of the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
period. P. moranensis var. neovolcanica, however, tends to grow on igneous rocks of the Eje Volcánico Transversal.
Madrean pine-oak woodlands
The Madrean pine-oak woodlands are subtropical woodlands found in the mountains of Mexico and the southwestern United States.The Madrean pine-oak woodlands are found at higher elevations in Mexico's major mountain ranges, the Sierra Madre Occidental, the Sierra Madre Oriental, the Trans-Mexican...
, or temperate montane woodlands. However, its distribution penetrates into tropical forests and xerophytic
Xerophyte
A xerophyte or xerophytic organism is a plant which has adapted to survive in an environment that lacks water, such as a desert. Xerophytic plants may have adapted shapes and forms or internal functions that reduce their water loss or store water during long periods of dryness...
shrublands, as well as in gorges and canyon walls with high environmental humidity
Humidity
Humidity is a term for the amount of water vapor in the air, and can refer to any one of several measurements of humidity. Formally, humid air is not "moist air" but a mixture of water vapor and other constituents of air, and humidity is defined in terms of the water content of this mixture,...
. P. moranensis prefers humid and shady environments, such as slopes by streams, gullies, or road cuts, or among leaf litter in sandy soil high in organic matter. Its ability to gather nutrients from the arthropod prey it catches allows it to grow in low-nutrient environments where other plants would usually out-compete it. As a result, it is often found in disturbed areas or on steep cliffs or hillsides. Since its roots do little more than provide anchorage, the plant requires little or no soil, and dense clusters can be found clinging onto boulders, moss or crags in rock faces, or even epiphytically
Epiphyte
An epiphyte is a plant that grows upon another plant non-parasitically or sometimes upon some other object , derives its moisture and nutrients from the air and rain and sometimes from debris accumulating around it, and is found in the temperate zone and in the...
on tree trunks. Common companion plants include moss
Moss
Mosses are small, soft plants that are typically 1–10 cm tall, though some species are much larger. They commonly grow close together in clumps or mats in damp or shady locations. They do not have flowers or seeds, and their simple leaves cover the thin wiry stems...
es, Selaginella, fern
Fern
A fern is any one of a group of about 12,000 species of plants belonging to the botanical group known as Pteridophyta. Unlike mosses, they have xylem and phloem . They have stems, leaves, and roots like other vascular plants...
s and other herbaceous plants, as well as canopy trees such as pines and oaks.
Cultivation
P. moranensis is one of the most popular and commonly cultivated Pinguicula, in part due to its large size, large and pretty flowers, and the ease with which it can be grown as a container plant. Most growers use an open soil mix composed of some combination of washed sandSand
Sand is a naturally occurring granular material composed of finely divided rock and mineral particles.The composition of sand is highly variable, depending on the local rock sources and conditions, but the most common constituent of sand in inland continental settings and non-tropical coastal...
, perlite
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...
, vermiculite
Vermiculite
Vermiculite is a natural mineral that expands with the application of heat. The expansion process is called exfoliation and it is routinely accomplished in purpose-designed commercial furnaces. Vermiculite is formed by weathering or hydrothermal alteration of biotite or phlogopite...
, peat moss, gypsum
Gypsum
Gypsum is a very soft sulfate mineral composed of calcium sulfate dihydrate, with the chemical formula CaSO4·2H2O. It is found in alabaster, a decorative stone used in Ancient Egypt. It is the second softest mineral on the Mohs Hardness Scale...
and/or decomposed granite. Soil should be kept well drained, but watered regularly with distilled water
Distilled water
Distilled water is water that has many of its impurities removed through distillation. Distillation involves boiling the water and then condensing the steam into a clean container.-History:...
in the summer and only very rarely once the plant enters its winter rosette. The species grows readily on well-lit windowsills, under fluorescent lights, or in warm to hot greenhouses.
Hybrids and cultivars
Although no natural hybrids involving P. moranensis have been reported, the species is known to hybridize quite readily in cultivation. As a result, a number of cultivarCultivar
A cultivar'Cultivar has two meanings as explained under Formal definition. When used in reference to a taxon, the word does not apply to an individual plant but to all those plants sharing the unique characteristics that define the cultivar. is a plant or group of plants selected for desirable...
s involving the species have been registered and are recognized by the International Carnivorous Plant Society
International Carnivorous Plant Society
The International Carnivorous Plant Society is a non-profit organization founded in 1972. It is the International Cultivar Registration Authority for carnivorous plants...
:
| cultivar name | parentage | description |
|---|---|---|
| Pinguicula 'George Sargent' Hort. Slack Adrian Slack Adrian Slack is an author and authority on carnivorous plants. He is the author of two books: Carnivorous Plants and Insect-Eating Plants and How to Grow Them .... |
P. moranensis × gypsicola Pinguicula gypsicola Pinguicula gypsicola is an insectivorous plant of the genus Pinguicula native to the Mexican state of San Luis Potosi, a heterophyllous member of the section Orcheosanthus. It grows in gypsum soils and forms stemless rosettes of upright, narrow leaves.-Morphology:P. gypsicola is a perennial... |
Lilac flowers, strap-shaped leaves, large winter rosettes. |
| Pinguicula 'Hameln' Hort. Studnicka | P. gypsicola Pinguicula gypsicola Pinguicula gypsicola is an insectivorous plant of the genus Pinguicula native to the Mexican state of San Luis Potosi, a heterophyllous member of the section Orcheosanthus. It grows in gypsum soils and forms stemless rosettes of upright, narrow leaves.-Morphology:P. gypsicola is a perennial... × moranensis |
Wider, P. moranensis like foliage. |
| Pinguicula 'John Rizzi' Hort. D'Amato Peter D'Amato Peter D'Amato is an American author, businessman, and carnivorous plant authority. He is the owner of California Carnivores, the largest nursery of carnivorous plants in the world, and the author of The Savage Garden , a book on the cultivation of insectivorous plants... |
P. moranensis × ? | Large, full flowers; oval, marginless undulating leaves. |
| Pinguicula 'L'Hautil' Hort. L.Legendre & S.Lavayssiere | P. (ehlersiae × moranensis) × moranensis | Two forms ("Grande" and "Petite"). P. 'Sethos' × P. 'Huahuapan' |
| Pinguicula 'Mitla' Hort. Studnicka | P. gypsicola Pinguicula gypsicola Pinguicula gypsicola is an insectivorous plant of the genus Pinguicula native to the Mexican state of San Luis Potosi, a heterophyllous member of the section Orcheosanthus. It grows in gypsum soils and forms stemless rosettes of upright, narrow leaves.-Morphology:P. gypsicola is a perennial... × moranensis |
Wider, P. moranensis like foliage. |
| Pinguicula 'Pirouette' Hort. J.Brittnacher, B.Meyers-Rice & L.Song | P. agnata Pinguicula agnata Pinguicula agnata is a tropical species of carnivorous plant in the family lentibulariaceae. Its flowers are a blue-violet color.... × (moranensis × ehlersiae) |
Hardy clone, attractive rosettes of pink leaves. |
| Pinguicula 'Sethos' Hort. Slack Adrian Slack Adrian Slack is an author and authority on carnivorous plants. He is the author of two books: Carnivorous Plants and Insect-Eating Plants and How to Grow Them .... |
P. ehlersiae × moranensis | Large flowers with a white star-like center. |
| Pinguicula 'Weser' Hort. Slack Adrian Slack Adrian Slack is an author and authority on carnivorous plants. He is the author of two books: Carnivorous Plants and Insect-Eating Plants and How to Grow Them .... |
P. moranensis × ehlersiae | Large flowers with single white streak down central lower lobe and dark veins. |
Additionally, three clones of P. moranensis have been registered as cultivars:
| cultivar name | origin | description |
|---|---|---|
| Pinguicula 'Huahuapan' Hort. Slack Adrian Slack Adrian Slack is an author and authority on carnivorous plants. He is the author of two books: Carnivorous Plants and Insect-Eating Plants and How to Grow Them .... |
Huajuapan de León Huajuapan de leon Heroica Ciudad de Huajuapan de León; in , meaning Place of Brave People) is a rural city with a surrounding municipality located in the northwestern part of the Mexican state of Oaxaca.... , Oaxaca Oaxaca Oaxaca , , officially Estado Libre y Soberano de Oaxaca is one of the 31 states which, along with the Federal District, comprise the 32 federative entities of Mexico. It is divided into 571 municipalities; of which 418 are governed by the system of customs and traditions... , Mexico |
Lilac-pink flowers with crimson touches at their base. |
| Pinguicula 'Libelulita' Hort. Rice & Salvia | southern Mexico | Square-tipped petals, pale purple at edges darkening to deep velvet-red near base, heavily veined, white center. |
| Pinguicula 'Vera Cruz' Hort. Slack Adrian Slack Adrian Slack is an author and authority on carnivorous plants. He is the author of two books: Carnivorous Plants and Insect-Eating Plants and How to Grow Them .... |
Veracruz Veracruz Veracruz, formally Veracruz de Ignacio de la Llave officially Estado Libre y Soberano de Veracruz de Ignacio de la Llave , is one of the 31 states that, along with the Federal District, comprise the 32 federative entities of Mexico. It is divided in 212 municipalities and its capital city is... , Mexico |
Deep rose with basal markings. |