Pinguicula orchidioides
Encyclopedia
Pinguicula orchidioides is a perennial rosette
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 5 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. Uniquely among Pinguicula species from the Americas, p. orchidioides produces gemma
Gemma (botany)
A gemma is a single cell, or a mass of cells, or a modified bud of tissue, that detaches from the parent and develops into a new individual. This type of asexual reproduction is referred to as fragmentation. It is a means of asexual propagation in plants...

-like basal buds which elongate into stolons and serve as a means of asexual reproduction
Asexual reproduction
Asexual reproduction is a mode of reproduction by which offspring arise from a single parent, and inherit the genes of that parent only, it is reproduction which does not involve meiosis, ploidy reduction, or fertilization. A more stringent definition is agamogenesis which is reproduction without...

. 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 purple flowers appear between July and September on upright stalks up to 22 centimeters long.

The species was first described in 1844 by Alphonse Pyrame de Candolle, but following an unfortunate misidentification by his contemporary William Jackson Hooker
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,...

, was relegated to the ranks of botanical synonymy and generally forgotten until it was rediscovered through the works of botanists in the 1990s.

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 orchidioides refers to dainty, orchid-like flowers.

Plant characteristics

Habit and Phenology

Like many Mexican butterworts, P. orchidioides 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 up to 10 centimeters (4 in) in diameter and composed of composed of ovate
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 lanceolate
Leaf shape
In botany, leaf shape is characterised with the following terms :* Acicular : Slender and pointed, needle-like* Acuminate : Tapering to a long point...

 leaves with distinct 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...

s. 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 November or December. 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 7–22 centimeters (3–9 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 from July-September as the summer growth begins.

Leaves and stolons

The leaf blades of the summer rosettes of P. orchidioides are smooth, rigid, and succulent, and generally green in color. The laminae
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 ovate
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 lanceolate
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 20 and 46 millimeters (2–5 in.) long and 6–18 millimeters wide, and have deeply involute
Involute
In the differential geometry of curves, an involute is a curve obtained from another given curve by attaching an imaginary taut string to the given curve and tracing its free end as it is wound onto that given curve; or in reverse, unwound. It is a roulette wherein the rolling curve is a straight...

 margins. These are supported by 10–30 millimeter 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...

s with ciliate margins.

The "winter" or "resting
Estivation
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. orchidioides is 6–13 millimeters (¼–½ in.) in diameter and consists of 25 to 36 small, compact, fleshy, non-glandular leaves. These are each 5 to 11 millimeters (–1 ¼ in.) long and one to three millimeters (–⅛ in.) wide, acuminate
Leaf shape
In botany, leaf shape is characterised with the following terms :* Acicular : Slender and pointed, needle-like* Acuminate : Tapering to a long point...

, acute, 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 :...

.

In a feature unique among Mexican Pinguicula, P. orchidioides produces stolons throughout the summer growing period. These start out as gemma-like buds in the winter rosette and elongate into whip-like stolons up to 8 centimeters (3 in) long during the summer. These stolons, which have small non-glandular leaves interspersed along their length, can take root to form new plantlets upon contact with a suitable growing substrate. This trait allows the species to form clumps of plants, many of which are genetically identical.

As is typical in the genus, the upper lamina surface of the summer leaves is 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. 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.

Flowers

P. orchidioides produces one to three flower
Flower
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 7 to 22 centimeters (3–9 in.) long, cherry red in color, and glabrous except near the calyx.

The flowers themselves are composed of five petal
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. Below the attachment point to the stem the petals are fused into a 18–26 millimeter long spur which protrudes backwards roughly perpendicular to the rest of the flower.

The violet-purple flowers are 30 to 48 millimeters (1 ¼–2 in.) long, and have a deeply bilabiate corolla, with an 2-lobed upper lip and a 3-lobed lower lip. The upper lobes are 10–14 millimeters (⅜–⅝ in.) long by 5–9 millimeters (–⅜ in.) wide and generally oblong-obovate. The outer lower lobes are oblong-lanceolate, 11–16 millimeters (¼–¾ in.) long by 4–7 millimeters (– in.) wide, narrowing at the tip. The central lower lobe is slightly longer than its neighbors, 15–19 millimeters (⅝–¾ in.) long by 4–5 millimeters (– in.) wide, and is marked with a white stripe at its base. The floral tube that houses the reproductive organs and is visible at the base of corolla lobes is short 3–4 millimeters (⅛– in.) long and lacks a palate.

The ovary and attached pistil protrude from the top of the floral tube near its opening, with the receptive stigma
Stigma (botany)
The stigma is the receptive tip of a carpel, or of several fused carpels, in the gynoecium of a flower. The stigma receives pollen at pollination and it is on the stigma that the pollen grain germinates. The stigma is adapted to catch and trap pollen with various hairs, flaps, or sculpturings...

 surface toward the front. Anthers hang from recurved 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 4–5 millimeter (– in.) dehiscent seed capsules containing numerous 1 millimeter long seeds.

Botanical history

P. orchidioides was first described by French-Swiss botanist Alphonse Louis Pierre Pyramus de Candolle in 1844 based on collections by G. Andrieux (130). De Candolle, in subdividing the genus Pinguicula
Pinguicula
The butterworts are a group of carnivorous plants comprising the genus Pinguicula. Members of this genus use sticky, glandular leaves to lure, trap, and digest insects in order to supplement the poor mineral nutrition they obtain from the environments. Of the roughly 80 currently known species, 12...

, included the species in the newly created section Orcheosanthus along with other species with purple, deeply bilabiate corollas with 5 sub-equal lobes, a short floral tube, and a large spur not protruding past this tube.

Two years later, Hooker
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,...

 described a plant he saw growing at Kew Botanical Gardens and, thinking it to be similar to Candolle's P. orchidioides, applied that name. Unfortunately, this specimen was actually a P. moranensis var. neovolcanica
Pinguicula moranensis
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 long, which are covered in mucilagenous glands that attract, trap, and digest arthropod...

, causing great confusion for taxonomists who thereafter treated P. orchidioides as a synonym of P. caudata or P. macrophylla.

Recognition of the species was only maintained by Sprague (1928), who contended that De Candolle's species was discrete from the plant described by Hooker. 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...

, while omitting the species from his revision of the genus in 1966, noted that P. orchidioides auct. non A.DC.: Hook.
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,...

was probably a nomen dubium vel ambiguum. However, the identification of P. orchidioides A.DC. was left unaddressed and was forgotten, so that when Hans Luhrs described stoloniferous Pinguicula specimens in 1995, he did so under a new name: P. stolonifera.

It wasn't until 1998 that the species was re-described under the name P. orchidioides by Mexican Pinguicula specialist Sergio Zamudio. He noted its distinction from other species based on leaf shape, stolon production, flower morphology and geographical isolation (from P. oblongiloba).

Distribution and habitat

P. orchidioides is currently known from only a handful of sites, most of them in the state of 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...

, though a few locations are also known from the state of 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....

 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 department
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 Sololá
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...

. The lack of records from 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...

 and other intermediary zones is likely the result of sampling bias. Here it grows in mountainous regions between 2000 and 2700 (3000) meters (6500–8850 ft) in altitude.

P. orchidioides seems to prefer humid hillsides, slopes or embankments, where it grows amongst pine-oak woodlands
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...

 in reddish-brown to brown, clay or sandy-clay soil. Other companion plants include members of the genera Arbutus
Arbutus
Arbutus is a genus of at least 14 species of flowering plants in the family Ericaceae, native to warm temperate regions of the Mediterranean, western Europe, and North America.-Description:...

, Arctostaphylos
Arctostaphylos
Arctostaphylos is a genus of plants comprised by the manzanitas and bearberries. They are shrubs or small trees.There are about 60 species of Arctostaphylos, ranging from ground-hugging arctic, coastal, and mountain species to small trees up to 6 m tall. Most are evergreen , with small oval...

, and Clethra
Clethra
Clethra is a genus of between 30-70 species of flowering shrubs or small trees. It is one of two genera in the family Clethraceae...

.

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