Pinguicula moranensis
Pinguicula moranensis | |
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In habitat in Oaxaca | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Eudicots |
Clade: | Asterids |
Order: | Lamiales |
Family: | Lentibulariaceae |
Genus: | Pinguicula |
Species: | P. moranensis
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Binomial name | |
Pinguicula moranensis | |
Varieties | |
P. moranensis var. moranensis | |
Synonyms | |
According to Zamudio (2001) unless otherwise noted.[1]
P. moranensis var. neovolcanica:
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Pinguicula moranensis 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
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
Plant characteristics
Habit
Pinguicula 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
Leaves and carnivory
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
As with all members of the genus, these leaf blades are densely covered by peduncular (stalked)
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.[13] 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.[8] These results suggest that the amount of investment in carnivorous features is an adaptation to the environmental gradients.
Winter rosette
The "winter" or "
Flowers
Pinguicula moranensis produces one to seven flowers 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 (1⁄8 in.) at the base to one millimeter (1⁄16 in.) at the top.[12]
The flowers themselves are composed of five
The ovary and attached
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 (1⁄4–3⁄8 in.) long by 4–9 millimeters (5⁄32–3⁄8 in.) wide and generally oblong, obovate, or cuneate. The lower lobes are similarly shaped and are 7–20 millimeters (1⁄4–3⁄4 in.) long by 4–18 millimeters (5⁄32–3⁄4 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 (5⁄32–1⁄4 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.[12] 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."[19] A rare white-flowered form is also known.[19]
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".[20] 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 Humboldt 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.[2] In 1817, Carl Sigismund Kunth described 3 new species from Alexander von Humboldt and Aimé Bonpland'sLatin American expedition: The Peruvian P. calyptrata and the first known Mexican species: P. macrophylla and P. moranensis.[21] At this point no infrageneric classification had yet been suggested.
In 1844, a French-Swiss botanist by the name of
The section Orcheosanthus grew as
Twentieth century
When Barnhart revised the family Lentibulariaceae 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.[24] 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.[6]
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.[5] Since that time 14 additional species have been discovered and assigned to the section.[25] When Zamudio redefined the section in 1999, however, he chose to include only 12 species, including all six of Casper's choices.[6] 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.
In further disagreement with Zamudio'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).[11] 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, Zamudio (1999) came to the conclusion that the species could be divided into two distinct
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.
Zamudio 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
Distribution and habitat
Pinguicula moranensis is the most widely distributed member of the
Pinguicula moranensis most often grows in oak,
Cultivation
Pinguicula moranensis is one of the most popular and commonly cultivated Pinguicula,
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 cultivars involving the species have been registered and are recognized by the International Carnivorous Plant Society:[28]
cultivar name | parentage | description |
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Pinguicula 'George Sargent' Hort. Slack | P. moranensis × gypsicola | Lilac flowers, strap-shaped leaves, large winter rosettes. |
Pinguicula 'Hameln' Hort. Studnicka | P. gypsicola × moranensis | Wider, P. moranensis like foliage. |
Pinguicula 'John Rizzi' Hort. D'Amato | 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[29] | P. gypsicola × moranensis | Wider, P. moranensis like foliage. |
Pinguicula 'Pirouette' Hort. J.Brittnacher, B.Meyers-Rice & L.Song | P. agnata × (moranensis × ehlersiae) | Hardy clone, attractive rosettes of pink leaves. |
Pinguicula 'Sethos' Hort. Slack | P. ehlersiae × moranensis | Large flowers with a white star-like center. |
Pinguicula 'Weser' Hort. Slack | 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:[2]
cultivar name | origin | description |
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Pinguicula 'Huahuapan' Hort. Slack | Huajuapan de León, Oaxaca, 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 | Veracruz, Mexico | Deep rose with basal markings. |
Notes
- ^ Zamudio 2001, p. 160–161, 182.
- ^ a b c d e f g Schlauer, Jan: Carnivorous Plant Database, version November 15, 2005, 16:25.
- ^ "Pinguicula moranensis Kunth". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 8 April 2023.
- ^ a b c d e f g Zamudio, S. 1999
- ^ a b Casper, S.J. 1966
- ^ a b c d e Zamudio, Sergio 2001[page needed]
- ^ As an example, see Speta 1989. Of the three species described by Speta & Fuchs, two (P. rectifolia and P. potosiensis) were rejected as synonyms by Zamudio 2001, while P. rectifolia was supported by Ciezlak et al.'s phylogenetic study in 2005.
- ^ a b c Alcalá, R.E. & Dominguez, C.A. 2005
- ^ Peduncle length: Zamudio 2001, p. 158.
- ^ According to Cieslak et al. 2005, P. moctezumae, P. moranensis, P. emarginata, P. species 'Huahuapan' and P. rectifolia all share this trait.
- ^ a b c d Cieslak et al., 2005
- ^ a b c d Zamudio 2001, p. 158.
- ^ a b Legendre, L. 2000
- ^ Zamudio 2001, p. 161.
- ^ According to Cieslak (2005), the spur length is a diagnostic feature when differentiating this species from P. recitoflia. Cieslak asserts that in P. moranensis the spur is more than 50% longer than the rest of the corolla.
- ^ a b c d e Zamudio 2001, p. 159.
- ^ Legendre, L. Pollination of Pinguicula flowers. Retrieved on January 2, 2007.
- ^ Zamudio 2001, p. 60.
- ^ a b Zamudio 2001, p. 184.
- ^ Zamudio 2001, p. 26; "La identidad de Pinguicula moranensis y su delimitación precisa es quizás el problema más difÍcil de resolver dentro del género."
- ^ From Zamudio 2001, orig. Humboldt, A., A. Bonpland & C.S. Kunth. 1817
- ^ From Zamudio 2001, orig. A.P. De Candolle 1844
- ^ Zamudio 2001, p. 26.
- ^ The difficulty of basing taxonomical work in the genus Pinguicula solely on dried specimens has been noted by several frustrated botanists. McVaugh & Mickel in their 1963 notes on the section Orcheosanthus noted that "Herbarium study of these plants is difficult and frustrating because the leaves and other vegetative and reproductive structures are so fragile that dried specimens are seldom adequately preserved. Usually but a single flower is found on a plant, and the extraordinarily fragile pressed flowers often lose their original color and shape to a very large extent. Often neither vegetative nor reproductive structures can be studied without severe damage to the specimen" (McVaugh, 1963). Barnhart (1916) had stated that "the range of variation is enormous, but how much is seasonal, how much individual, and how much of taxonomic importance, is (based on herbarium samples alone) at present mere guesswork" (Barnhart, 1916). This conclusion was echoed by Sprague (1928), Ernst (1961), and McVaugh (1963).
- ^ Zamudio 2001, p. 29.
- A. De Candolle. Hooker described a plant he saw growing at Kew Botanical Gardens and, thinking it to be similar to Candolle's P. orchidioides, applied that name.
- ^ a b D'Amato P. 1998, p. 203–205.
- ^ International Carnivorous Plant Society, Registered Cultivar Names—Pinguicula Archived 2007-03-18 at the Wayback Machine. Retrieved on January 2, 2007.
- ^ An illegitimate homonym of this cultivar exists. Peter D'Amato describes a P. moranensis clone under this name in The Savage Garden (1998).
References
- Alcalá, R.E. & Dominguez, C.A. 2005 Differential Selection for Carnivory Traits Along an Environmental Gradient in Pinguicula moranensis; Ecology, 86(10), 2005, pp. 2652–2660
- Casper, S.J. 1966. Monographie der Gattung Pinguicula L. Heft 127/128, Vol 31. Stuttgart: Bibliotheca Botanica
- Cieslak, T., Polepalli, J.S., 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; American Journal of Botany. 2005;92:1723–1736.
- Candonelle, A.P. De. 1844. Prodromus systematis naturalis regni vegetabilis; Tomo VIII. Paris. pp. 26–32.
- D'Amato, P. 1998. ISBN 0-89815-915-6
- Ernst, A. 1961 Revision der Gattung Pinguicula Bot. Jahrb. Syst. 80(2): 145–194
- Hemsley, W.B. 1879–1888. Botany. In: Godwin, F.D. & O. Salvin, Biologia Centrali-Americana. R.H.Porter, London. 5 vol.
- Hooker, J.D. 1846 P. orchidioides. Botanical Magazine. 72: tab. 4231
- Humboldt, A., A. Bonpland & C.S. Kunth. 1817. Nova genera et species plantarum. II: pp. 225–226
- Legendre L (2000). "The genus Pinguicula L. (Lentibulariaceae): an overview". Acta Botanica Gallica 141 (1): 77–95.
- Mc Vaugh, R. & J.T.Mickel 1963. Notes on Pinguicula, sec. Orcheosanthus. Brittonia, Vol. 15, Issue 2 (apr. 15, 1963), p. 134–140
- Morren, E. 1872. Notice sure le Pinguicula flos-Mulionis Ed. Mn. ou Grassette Fleur de Muletier, Fig. Pl. XXVII. Belgique Hort. 22:371–374
- Zamudio Ruiz, S. 2001 Revision de la seccion Orcheosanthus del Genero Pinguicula (Lentibulariaceae) Universidad Nacional Autónoma de México.
- Sessé, M. & J. M. Mociño. 1893. Plantae Novae Hispaniae. 2a. Ed. México. Oficina Tipográfica de la Secretaria de Fomento, México, D.F. 175 pp.
- Speta, F. & F. Fuchs 1989 Drei neue Pinguicula-Arten der Sektion Orcheosanthus DC. aus Mexiko; Phyton (Austria), vol. 29, fasc. 1, pp. 93–103
- Sprague, T.A. 1928. The orchid-flowered butterworts. Kew Bull. 6: 230–234
- Watson, W. 1911. Pinguicula rosei. Gard. Chron. ser. 3, 29: 82.
- Zamudio, S. 1999 Notas sobre la identidad de Pinguicula moranensis H.B.K., con la description de una variedad nueva, Acta Botanica Mexicana, 1999, vol.49, pages 23–34
External links
- A World of Pinguicula — Articles, photos, taxonomic and horticultural information
- Carnivorous Plant Photo Finder: Pinguicula
- International Carnivorous Plant Society
- Pinguicula Discussion Forum