Utricularia resupinata | |
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1913 botanical illustration. | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Eudicots |
Clade: | Asterids |
Order: | Lamiales |
Family: | Lentibulariaceae |
Genus: | Utricularia |
Subgenus: | Utricularia subg. Utricularia |
Section: | Utricularia sect. Lecticula |
Species: | U. resupinata |
Binomial name | |
Utricularia resupinata Greene ex Bigelow | |
Synonyms | |
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Utricularia resupinata, popularly known as lavender bladderwort or northeastern bladderwort, [1] [2] is a small perennial subaquatic carnivorous plant that belongs to the genus Utricularia (family Lentibulariaceae ). It is native to eastern Canada, the United States, and Central America. This plant species has an interesting etymology, growth pattern, ecology, and research history. As a plant that is threatened or endangered in many of the States where it is found, it is a candidate for sound conservation efforts.
Botanical publications of this species will often describe it as "Utricularia resupinata B. D. Greene ex Bigelow," identifying its scientific name (Utricularia resupinata), its genus (Utricularia), its specific epithet (resupinata), the original discoverer (Benjamin Daniel Greene), and the publisher of the first information about the species (Jacob Bigelow). Scientific names in botany are defined by what is called a plant's binomial nomenclature or the use of two names for a species, genus and epithet. The person who describes and publishes a new species cannot name it for him or herself. A plant may be named after the person who first collected it, or for a location, a plant feature, much more. In this case, the epithet is derived from the Latin word resupinata. Etymology for Utricularia resupinata comes from two Latin words – utriculus which means "a small bottle" and refers to its insect-trapping bladders, and the word resupinata which means "bent or thrown back", for the top part of the bladderwort's flower. [3]
The 1913 botanical illustration of this bladderwort species depicts a delicate 2 to 12-inch stem growing along or just below the surface in very shallow water on a slender root or basal system; leaves are tiny or absent, often buried in the sand or mud; the showy blue to purple flower blooms from August to September with a two-lipped petal held up by a thin stem, the upper lips facing upwards and the three-lobed lower lip having a projection or sac extending from the petal base; fruit forms in a two-valved sac holding small seeds, on a separate stem emerging from a bract just above the base of the plant, as if its stem is sitting on a couch or in a flower pot, the fruit being dry and splitting open when ripe; and bladderwort reproduces both sexually by seed and asexually by producing compact buds or turions which break free from the parent plant and spread out nearby to start new plants. Native Plant Trust's "Go Botany" lists Utricularia resupinata's characteristics in minute detail, stating that the plant has a lifespan of two-years or more. [4] The attribute of this carnivorous plant for capturing and digesting animal life is the main factor placing it in the Lentibulariaceae or bladderwort family – with a typical plant system holding from a dozen to a hundred 1 to 6 mm bladders or "traps" for minute animal life. [5] The flower pictured here is a good illustration of this bladderwort's showy lavender flower with its lower projecting sac. [6]
Utricularia resupinata grows on the edges of wetlands or along the shore or in the shallow water of ponds, lakes, or rivers. It can be found in the moist sandy soil of recently built roads. Ideal growing conditions consist of a sandy substrate covered over by a thin layer of mud or muck. In its northern range it appears to only flower when low water levels occur at the same time as higher than average temperatures exist. [7] It was thought to be extirpated in Indiana until it was rediscovered in 2005. [2]
The specimen of Utricularia resupinata depicted below was collected by George R. Cooley, R. J. Easton, Carroll E. Wood, Jr., and C. Earle Smith, Jr. in May, 1961 on the shore of Lake Tsala Apopka, Florida, in a half inch of water. This specimen is part of the University of South Florida herbarium, and it shows this specimen in the flowering stage, with its thick network of stems and base runners, dried flowers and tiny leaves and bladders, and stems with seeds. [8]
Professor Asa Gray published the first of eight editions of his Manual of Botany in 1848, with a limited range noted for collected specimens of Utricularia resupinata, "at sandy margins of ponds, East Maine to Rhode Island." [9] The "Bulletin of the Torrey Botanical Society" – published in New York City by the first such club in the Americas – was quick to announce that a member of the Syracuse Botanical Club had found it beyond this limited zone "in the North Woods, Fenton's No. 4, Lewis Co., New York, Aug., 1879, on the marshy shores of a lake, as we are informed." [10]
Frank Tweedy had been hired in 1876 to survey and map the Beaver River basin for Verplanck Colvin's Adirondack Survey in New York (state), pursuing his passion of plant collecting each Sunday, and he responded to that announcement in the Torrey Bulletin with his own discovery of Utricularia resupinata in several more locations. This was the first of the more than 6,000 officially reported specimens Tweedy collected, most of them in the Rocky Mountains:
I collected that plant in August, 1875, at the same locality…namely, muddy shore of Beaver Lake, No. 4, Lewis Co., N.Y. During the past season I collected quite a number of the same plant on the shore of Big Moose Lake, Herkimer Co., N.Y., and at Twitchell Lake, in the same county. I do not think it is uncommon through Northern New York. [11]
Since Tweedy's finds deep in the Adirondack Mountains, Utricularia resupinata has been found in Canada, the eastern US as far as the Great Lakes states, and in Central America. A Range Map created by "The Floristic Synthesis of North America (BONAP)" shows the wider range for this species, though it is increasingly threatened or endangered in many States. [12]
At the top of the taxonomy or scientific classification offered here is the kingdom of plants (Plantae), followed by a number of "clades" or monophyletic groups composed of a common ancestor and all its lineal descendants: The clade of vascular plants (Tracheophytes); the clade of flowering plants (Angiosperms); the clade of flowering plants with two seeds on germination (Eudicots); the clade of flowering plants with only one common ancestor (Asterids); and the order of flowering plants as a subgroup of the Asterids with specific characteristics such as opposite leaves (Lamiales). This huge order of Lamiales includes 23,810 species, 1,059 genera, and is divided into 24 families of plants. [13]
Utricularia resupinata fits into all of these larger plant categories, with its family (biology)—the next category in the scientific classification—including only carnivorous plants (Lentibulariaceae). This family is made up of three genera—corkscrew plants (Genlisea), the butterworts (Pinguicula), and the bladderworts (Utricularia). Within this last group or genus of 240 species we find Utricularia resupinata, with its "synonyms" or taxon of plants that experienced a name change. Listed here is Lecticular resupinata, the name given to Ultricularia resupinata by Barnhart in 1913 and Utricularia greenei, the name given it by Oakes in 1841. Lecticular is Latin for "couch" or "cubicle" and was chosen because of the unique bract on this plant's lower stem. [14]
The Angiosperm Phylogeny Group (APG) -- named for the clade of flowering plants—is an international group of botanists who are working to establish a standard plant taxonomy for all flowering plants. Earlier taxonomies tended to be set by a given nation or botanical school, hence a multitude of differences. The first APG system for categorizing plants was set up in 1998 with updates in 2003 (APG II), 2009 (APG III), and 2016 (APG IV). [15] The scientific classification or taxonomy offered in this article is based on the more recent work of APG III and IV.
The list offered on the US Department of Agriculture's website reveals just how immense the earth's "Tree of Life" really is, and this eight-page list covers just the main divisions, families, orders, and classes for flowering plants in the much larger kingdom of plants. [16] A search of this listing finds the kingdom, order, and family for Utricularia resupinata.
"Utricularia resupinata B.D. Greene ex Bigelow" identifies the species' scientific name with the first person to discover this flowering plant—a Massachusetts lawyer named Benjamin Daniel Greene (1793–1862), whose finding was reported in what became the standard botanical resource for that time period, with the following brief note: "Greene, M.S. Greene's bladder wort…A small delicate species with purple flowers, discovered by B. D. Greene, Esq. at [a pond in] Tewksbury. Middlesex Co, MA." [17] : 10 American botanist, physician, and botanical illustrator Jacob Bigelow (1787–1879) published his first edition of Florula Bostoniensis in 1814 – a detailed survey of flora in the greater Boston region, adding neighboring New England states in later editions.
Two of Frank Tweedy's bladderwort finds are currently in New England herbaria, one at Harvard University and the other at the University of Vermont, both collected in 1879 on the shore of Big Moose Lake, Herkimer County, NY. [18] A herbarium (plural herbaria) is a collection of plant specimens that have been preserved for scientific study. [19] A search of all herbaria for this species in the Mid-Atlantic Herbaria Consortium yields 609 finds, beginning with B.D. Greene's specimen now displayed in the New York Botanical Garden, included on this list with his date of discovery, 1829. The rest of the collection ranges from then to the most recent specimen found at Lake Hartwell, S.C. on October 9, 2019, with this description: "A Colony growing en masse, on bank in area where lake waters have receded. Proliferous colony stretching far up the shore and in the turn of the cove. Leaves matted in mud, some plants producing early fruits. Flowers very pale pink." [20]
One of the early students of Utricularia resupinata and its carnivorous family ( Lentibulariaceae ) was Mary Treat (1830-1923), a naturalist who made major contributions in botany and entomology. She spent many hours over her microscope observing the bladders or traps along its supporting system of stems, trying to discover how "these little bladders" trapped and digested their animal prey. It was originally believed that the series of bladders on the stems and roots "floated the plant." Mary Treat was "one of the first scientists to suspect that the bladders were actually traps for tiny creatures rather than air floatation devices." [21]
In a book titled Through a Microscope, Mary Treat contributed a whole chapter on the Utricularia genus, puzzling over the "wonderful" process by which these bladders worked:
I have found almost every swimming animalculæ with which I am acquainted, caught in these vegetable traps; and when caught they never escape. Their entrance is easy enough; there is a sensitive valve at the mouth of the bladder, which, if they touch it, flies open and draws them in as quick as a flash. These downward-opening bladders not only entrap animalculæ, but, more wonderful still, the strong larvæ of insects. [22] : 62–63
Mary also held a five-year correspondence with biologist Charles Darwin as he was researching carnivorous plants, debating the question of how insects entered these bladders, finally convincing him of her theory. She recounted this in the same chapter, and it is worth quoting at length for this back story from the early history of botany:
Those who have read Mr. Darwin's very interesting book on Insectivorous Plants, will have noticed that he says the valve of Utricularia is not in the least sensitive, and that the little creatures force their way into the bladders -- their heads acting like a wedge. But this is not the case, as Mr. Darwin himself was convinced some years before his death. In his usual kind, gracious manner he admitted that he was wrong, and gracefully says the valve must be sensitive, although he could never excite any movement. In a letter to me bearing date June 1st, 1875, he says: "I have read your article (in Harper's Magazine) with the greatest interest. It certainly appears from your excellent observations that the valve is sensitive...I cannot understand why I could never with all my pains excite any movement. It is pretty clear I am quite wrong about the head acting like a wedge. The indraught of the living larva is astonishing." [22] : 69
Utricularia resupinata was moved to a new genus as the science of botany progressed, and New York Botanical Garden (NYBG) leader John Hendley Barnhart (1871-1949) categorized this species in 1913 under "Lecticula", a section of the genus Ultricularia. He based this taxonomy change on its unusual stem bracts noted above: "Utricularia sect. Lecticula is a section in the genus Utricularia that was originally described as genus Lecticula in 1913 by John Hendley Barnhart. The two species in this section are small subaquatic carnivorous plants that are distinguished by the unique bracts, which are basifixed and tubular. Both species are native to North and South America." [23] Botanist Norman Taylor (1900-1975) in association with the NYBG published Flora in the Vicinity of New York in 1915, reporting on Barnhart's new category (and name) for Utricularia resupinata. Under the Lentibulariaceae or bladderwort family of plants, he identified three genera -- Vesiculina Raf., Utricularia L., and Lecticula Barnhart, with Utricularia resupinata printed with its new name in this third genus. The following comment described this section of the genus: "L. [Lecticula] resupinata (B.D. Greene) Barnhart. In sandy bogs and borders of ponds: Me. to Fla., west to Mish. Rare." [24] Lecticula resupinata is thus a plant synonym for Utricularia resupinata.
Peter Geoffrey Taylor (1926-2011) put in 41 years of research and observation on the genus Utricularia. In 1989 he published The Genus Utricularia: A Taxonomic Monograph with Utricularia resupinata one of the 240 species in this genus, and now under its original name. Taylor illustrated 214 of these species in his book, considered to be ground breaking with its in-depth study of an entire genus: "Taylor's species list and classification are now generally accepted with some additions of newly described taxa and modifications based on phylogenetic studies. [25]
Mary Treat repeatedly observed the tiny bladders of the Utricularia genus under her microscope trap minute animal life as a mosquito larva, for example, triggered sensitive filaments at the mouth of the bladder, snapping it shut. But one mystery baffled her: "I soon became satisfied that the valve was very sensitive when touched at the right point, but to this day I cannot tell what the power is that so quickly draws the creatures within." [22] : 68–69 Recent research based on phylogenetics has answered some questions and raised others. The Utricularia bladder has long fascinated scientists. "Although the Utricularia traps are the smallest among those of carnivorous plants, they are arguably the most sophisticated and intricate ones." [5] : 640
Czech Republic botanist Lubomir Adamec has summarized the extensive research that has recently been done with species in the Utricularia genus – with a focus on its carnivorous bladder. Three things are clear. This plant expends a tremendous output of energy through its string of bladders. "When a prey species touches sensory hairs situated on the trap door it opens, the small animal is aspirated into the trap and the door closes again. This process of firing is complete within 10 -- 15 ms [milliseconds] and is the most rapid plant movement known." [5] The source of this ATP energy -- Adenosine triphosphate, the molecule for storing and transferring energy in cells—is still an open research question. This Utricularia resupinata picture of two bladders on the plant's stem captures the luminescence of these bladders with their "trap door" and trigger filaments branching out. [26]
Second, not all of the minute animals sucked into the Utricularia traps are digested, because it has been found that some of the organisms in this "bladder soup" actually assist in prey decomposition, a process similar to what happens in an efficient septic system. "In spite of its tiny volume, the trap fluid in Utricularia plants is inhabited permanently by various commensal microorganisms -- bacteria, cyanobacteria, microfungi, algae, euglena, dinophytes, protozoa (ciliates) and rotifers -- which live in a mutualistic interaction with the plant." [5] This "miniature food web" offers many ecological possibilities, with potential applications pending further research.
And third, the presence or absence of oxygen inside the bladder is a key part of this "sophisticated and intricate" process. "Therefore, captured organisms either die of O2 deprivation and are prey, or are able to tolerate anoxia and are commensals. Utricularia traps likely kill their prey by suffocation." [5] Lubomir Adamec's research summary while technical, invites the reader to draw on a knowledge of all the physical and life sciences. [5] An intricate collection of valves inside the bladder help govern this complex organ—inviting the reader to appreciate its mechanics, the pumping of water in and out at high speed and pressure; its electro-chemistry, transferring enough voltage when triggered to perpetuate the process; and learning of the symbiosis of minute animal species inside this botanical bladder, either facilitating or becoming food for digestion.
There are substantial concerns over the conservation status of Utricularia resupinata throughout its geographical distribution, with the following list offered by the US Department of Agriculture's source on "Threatened and Endangered Information:" "Of Special Concern" are Rhode Island and Tennessee; listed as "Threatened" are Massachusetts and Vermont; "Endangered" include Connecticut, Maine, Maryland, and New Jersey; and labeled as "Extirpated" are Indiana and Pennsylvania, with a recent discovery of a species community in the former. [27] Three reasons are offered for this species being of special concern, threatened, endangered, or extirpated—heavy recreational use of waterways; high nutrient levels from lawn fertilizers or faulty septic systems; and competition from native or non-native invasive species. [28] A conservation plan is needed for the protection and proliferation of this delicate but important species which so fascinated Frank Tweedy, Mary Treat, Charles Darwin, and generations of botanists and biologists.
Droseraceae is a family of carnivorous flowering plants, also known as the sundew family. It consists of approximately 180 species in three extant genera. Representatives of the Droseraceae are found on all continents except Antarctica.
Lentibulariaceae is a family of carnivorous plants containing three genera: Genlisea, the corkscrew plants; Pinguicula, the butterworts; and Utricularia, the bladderworts.
Utricularia, commonly and collectively called the bladderworts, is a genus of carnivorous plants consisting of approximately 233 species. They occur in fresh water and wet soil as terrestrial or aquatic species across every continent except Antarctica. Utricularia are cultivated for their flowers, which are often compared with those of snapdragons and orchids, especially amongst carnivorous plant enthusiasts.
Genlisea is a genus of carnivorous plants also known as corkscrew plants. The 30 or so species grow in wet terrestrial to semi-aquatic environments distributed throughout Africa and Central and South America. The plants use highly modified underground leaves to attract, trap and digest minute microfauna, particularly protozoans. Although suggested a century earlier by Charles Darwin, carnivory in the genus was not proven until 1998.
Blandfordia, commonly known as Christmas bells, is a genus of four species of flowering plants native to eastern Australia. Christmas bells are tufted, perennial herbs with narrow, linear leaves and up to twenty large, drooping, cylindrical or bell-shaped flowers.
Haptanthus is a monotypic genus containing the sole species Haptanthus hazlettii, a shrub or small tree known only from the locality of Matarras in the Arizona Municipality in Honduras. Its flowers are unique among the flowering plants. A single "female" (carpellate) flower has two branches on either side which carry "male" (staminate) flowers. The flowers are very simple, lacking obvious sepals or petals. The family placement of the genus has been uncertain, but based on molecular phylogenetic research, it is included in the family Buxaceae as of September 2014. Very few individuals have ever been found and its habitat is threatened by logging.
Genlisea margaretae is a carnivorous species in the genus Genlisea native to areas of Madagascar, Tanzania, and Zambia. It has pale bundles of root-like organs up to about 20 cm long under ground that attract, trap, and digest protozoans. These organs are subterranean leaves, which lack chlorophyll. It had been known to possess the smallest known genome of any flowering plant as of 2006, but was later surpassed by the related species Genlisea tuberosa.
Utricularia dichotoma, commonly known as fairy aprons, is a variable, perennial species of terrestrial bladderwort. It is a widespread species with mauve or purple fan-shaped flowers on a slender stalk and usually grows in wet locations.
Utricularia gibba, commonly known as the humped bladderwort or floating bladderwort, is a small, mat-forming species of carnivorous aquatic bladderwort. It is found on all continents except Antarctica.
Philcoxia is a genus of seven rare plant species in the Plantaginaceae that are endemic to Brazil and resemble terrestrial species of the genus Utricularia. The genus, formally described in 2000, consists of the species P. bahiensis, P. goiasensis, P. minensis, P. tuberosa, P. rhizomatosa, P. maranhensis and P. courensis, each of the first three named for the Brazilian state to which it is endemic. The species are characterized by subterranean stems, peltate leaves at or below the soil surface, and five-lobed calyces. Their habitat has been reported as areas of white sand in the midst of cerrado vegetation at an elevation between 800 and 1450 m. Initial descriptions of the genus included suspicions that the plethora of stalked capitate glands on the upper surfaces of leaves was an indication that these species may be carnivorous. A study published in 2007 tested P. minensis for protease activity, a typical test for the carnivorous syndrome, and could detect none. Later studies detected other digestive enzymes such as phosphatases and qualitatively assessed prey digestion and nutrient uptake, suggesting that it is a true carnivorous plant. The genus epithet honors David Philcox (1926-2003), a botanist at Kew Gardens who worked extensively in tropical Scrophulariaceae.
Utricularia inflata, commonly known as the swollen bladderwort, inflated bladderwort, or large floating bladderwort, is a large suspended aquatic carnivorous plant that belongs to the genus Utricularia. It is a perennial that is native to the southeastern coastal plains of the United States. It has often been confused with U. radiata, which is similar but smaller than U. inflata. Since 1980, U. inflata has been reported to exist in locations beyond its traditional range, such as the Adirondack Mountains in New York, southeastern Massachusetts, and in Washington State. Studies on the populations in the Adirondacks suggest that an introduction of U. inflata to a location where it naturalizes can lead to altered sediment chemistry by reducing the net primary productivity of native species. It is also listed by the state of Washington as a problematic species because of the dense mat-forming habit of this aquatic Utricularia. It is one of the few carnivorous plants that can be invasive.
Utricularia macrorhiza, the common bladderwort, is a perennial suspended aquatic carnivorous plant that belongs to the genus Utricularia. U. macrorhiza is native to North America and eastern temperate Asia.
Utricularia purpurea, the eastern purple bladderwort, is a medium-sized suspended aquatic carnivorous plant that belongs to the genus Utricularia. U. purpurea is endemic to North and Central America. It has been suggested that U. purpurea may have partially lost its appetite for carnivory. Richards (2001) did an extensive study in the field on it and noted that trapping rates of the usual Utricularia prey were significantly lower than in other species in the genus. Richards concludes that this species can still trap and digest arthropod prey in its specialized bladder traps, but does so sparingly. Instead, it harbors a community of algae, zooplankton, and debris in the bladders that indicates U. purpurea favors a mutualistic interaction in place of a predator–prey relationship.
Peter Geoffrey Taylor (1926–2011) was a British botanist who worked at Royal Botanic Gardens, Kew throughout his career in botany. Taylor was born in 1926 and joined the staff of the herbarium at Kew in 1948. He published his first new species, Utricularia pentadactyla, in 1954. In 1973, Taylor was appointed curator of the orchid division of the herbarium and, according to Kew, "under his direction, orchid taxonomy was revitalised and its horticultural contacts strengthened."
Utricularia livida, the leaden bladderwort, is a species of flowering plant in the bladderwort family, native to central and southern Africa, and Mexico. Growing to 50 cm (20 in) tall and broad, it is a carnivorous perennial. It was originally described and published by Ernst Heinrich Friedrich Meyer in 1837.
Utricularia sandersonii, Sanderson's bladderwort, is a species of flowering plant in the bladderwort family. Originally described and published by the British botanist Daniel Oliver in 1865, it is a carnivorous evergreen perennial, endemic to northern KwaZulu-Natal and Transkei in South Africa.
Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans, typically insects and other arthropods, and occasionally small mammals and birds. They still generate all of their energy from photosynthesis. They have adapted to grow in waterlogged sunny places where the soil is thin or poor in nutrients, especially nitrogen, such as acidic bogs. They can be found on all continents except Antarctica, as well as many Pacific islands. In 1875, Charles Darwin published Insectivorous Plants, the first treatise to recognize the significance of carnivory in plants, describing years of painstaking research.
The Genus Utricularia: A Taxonomic Monograph is a monograph by Peter Taylor on the carnivorous plant genus Utricularia, the bladderworts. It was published in 1989 by Her Majesty's Stationery Office (HMSO) as the fourteenth entry in the Kew Bulletin Additional Series. It was reprinted for The Royal Botanic Gardens, Kew in 1994.
Genlisea tuberosa is a carnivorous species in the genus Genlisea that is endemic to Brazil and found only in campos rupestres vegetation. Lacking any roots, it has unpigmented bundles of "rootlike" subterranean organs, technically leaves, which attract, trap, and digest protozoans. This species is unique in the genus in its formation of tubers. As of 2014, Genlisea tuberosa has the smallest known genome of any flowering plant, at 61 Mbp, or 61,000,000 base pairs.
Desmodium tweedyi is an herbaceous flowering plant in the pea family native to northern Texas and southern Oklahoma popularly known as "Tweedy's ticktrefoil" or "tick-clover." The legume or seed pod it produces has given the species its common names from its ability to cling to clothing. Along with other species in the Desmodium genus, D. tweedyi has become a candidate for soil enrichment, suppression of insect pests, mulch and green manure production, and making "good fodder for animals including bobwhite, turkey, grouse, deer, cattle and goats."
Barry Rice captured this picture of Lavender Bladderwort flower, scientific name Utricularia resupinate in 2010
This specimen is Catalog #43122
This list includes classifications and principal scholarly source for each, including a main outline of a fossil record for the plant kingdom; this web page includes a link to the "Tree of Life" for the Green Plant BAC Library Project maintained by the Arizona Genomics Institute.
Entering the species scientific name and the collector Frank Tweedy, brings up his two finds; entering just the scientific name reports all specimen finds.
A leading botanical collection of approximately 3 million specimens, representing half to two thirds of the world's flora
This specimen was collected by D.Z. Damrel and K.J. Alford
Plant Profile Utricularia resupinata, Reversed Bladderwort
This picture of the Utricularia resupinate bladders or traps was taken by Peter M. Dziuk in 2015