Bifrenaria
Encyclopedia
Bifrenaria, abbreviated Bif. in horticultural trade, is a genus of plant
in family Orchidaceae
. It contains 20 species found in Panama
, Trinidad
and South America
. There are no known uses for them, but their abundant, and at first glance artificial, flower
s, make them favorites of orchid growers.
The genus can be split in two clearly distinct groups: one of highly robust plants with large flowers, that encompass the first species to be classified under the genus Bifrenaria; other of more delicate plants with smaller flowers occasionally classified as Stenocoryne or Adipe. There are two additional species that are normally classified as Bifrenaria, but which molecular analysis indicate to belong to different orchid groups entirely. One is Bifrenaria grandis which is endemic to Bolívia
and which is now placed in Lacaena
, and Bifrenaria steyermarkii
, an inhabitant of the northern Amazon Forest, which does not have an alternative classification.
Bifrenaria are generally robust plants, of sympodial
growth, between ten and sixty centimeters tall. They are characterized by round-section root with thick velamen
, four-angled fleshy pseudobulb
s of one internode, often basally protected by dried sheaths and with only one apical leaf
(except for Bifrenaria steyermarkii, which occasionally has two), plicate (fan-folded) enervated leathery leaves, yet malleable and not exceedingly thick, with a pseudo-petiole of basal round section, and a basal inflorescence
s bearing up to ten flowers, which seldom surpass the leaves' length.
Bifrenaria flowers are strongly scented, they have sepals slightly larger than the petals, with the lateral ones basally united to the column
foot forming a calcar
with truncated extremity.The column is slightly arching, generally without wings or any other appendages, bearing a foot which the labellum is hinged to, whose shape varies, articulated to the column, with a longitudinal channeled callus often with a basal claw. Flowers show two elongated stipe
s, hardy ever one, at least twice longer than wide, with salient viscidium, visible caudicles and retinacle in inverted positions. The superposed pollinia number four, and are protected by a deciduous incumbent anther. Fruits are green, erect or pendulous; they take about eight months to ripe and hold hundreds of thousand yellowish or brownish elongated seeds up to 0.35 mm long. Among all the mentioned, the main characteristic distinguishing Bifrenaria from its closest relatives is the presence of the calcar. Other important characters are the four-sided single-leaved pseudobulbs besides the raceme inflorescence with two to ten flowers.
Little is known about pollination in Bifrenaria. Apparently the only existing records report the presence of some large species' pollinia observed on the back of male Eufriesea violacea bees (Euglossinae), and of Bombus brasiliensis (Bombini). Although there are no reports of flower pollination being directly observed, a paper published in 2006 studied the micromorphology of the labellum in Bifrenaria species, looking for substances useful to insect
s as food. The absence of such substances on the densely pubescent surface of most Bifrenaria labelli seems to indicate possible pollination by large bees as the major mean. Another indicator of this possibility is the strong smell emanated by species like B. tetragona which are similar to those of plants in other families which are also pollinated by these bees. The smaller pubescent species may be pollinated by smaller bees, while the smooth ones, which have strong colored flowers, as B. aureofulva, might be pollinated by hummingbird
s.
, under the name Dendrobium harrisoniae
. Three years later, he also described the first small-flowered species, B. racemosa, but placed it in Maxillaria
. With these two publications began a long series of species descriptions and confusing genera creation which generated much doubt for the next two centuries. The Royal Botanic Garden registers the submission of 69 species or subspecific taxa under Bifrenaria since the description of the first species. Among these, twenty are generally accepted but only seventeen are truly well established, with no doubts about their limits and classification. Thirteen other species are still accepted but now placed in other genera, and four or five, due to deficiencies in their descriptions, might never be definitely identified.
In 1832, John Lindley
proposed the genus Bifrenaria and described its type species
, Bifrenaria atropurpurea
, previously named by Conrad Loddiges as Maxillaria atropurpurea. The name Bifrenaria comes from bi, two, and freno, brake, a reference to the shape of the two pairs of pollinia hold by separated caudicles presented by its flowers.
In 1837, Constantine Samuel Rafinesque, considering the noticeable vegetative difference between the few Bifrenaria known at the time, proposed the genus Adipe, based on B. racemosa morphology, described by Hooker few years before, to which he added the description of a supposed new species, Adipe fulva (today treated as a synonym of B. racemosa). The next year Lindley received a specimen from Amazonia, which was morphologically even more distant from the known species, but nonetheless described it as Bifrenaria longicornis
. Five years later, apparently not aware of Rafinesque's previous genus Adipe, Lindley changed his mind and suggested that this species should be classified under a new genus, Stenocoryne. Six species were subsequently attributed to Stenocoryne by various taxonomists, but the genus Rafinesque proposed remained unused until 1990.
Two species similar to Bifrenaria, but that showed a highly salient claw at the base of the labellum and lateral lobes abruptly divided were then classified under this genus. In 1914, Rudolf Schlechter suggested they should be classified under the genus Lindleyella, with Lindley's Bifrenaria aurantiaca
(which presented the mentioned differences) as the type. However this genus name was already occupied (It is a synonym of Lindleya
, in the Rosaceae
). Just thirty years later, in 1944, Frederico Carlos Hoehne, working on the first revision of genus Bifrenaria, corrected the suggestion of Schlechter. Hoehne initially proposed the genus Schlechterella for these species but, coincidentally, this name was also taken, this time by African Asclepiadaceae
. Finally a genus with an available name in homage to Schlechter was erected in the very next issue of the journal, Rudolfiella
, by which time time the number of species had increased to seven. On this revision, besides Rudolfiella, Hoehne divided Bifrenaria into two genera, accepting Lindley's Stenocoryne but calling attention to the existence of Rafinesque's Adipe, which should have nomenclatural priority, while also raising doubts about the identity of several described species. In 1990, Manfred Wolff formally resurrected the genus Adipe and transferred to it ten Bifrenaria species, besides the two already described by Rafinesque; his change was purely nomenclatural and he did not revisit the species.
Making the picture even more complex, in 1994, Karheinz Senghas, based on several characteristics shared only by B. tetragona and B. wittigii, described the genus Cydoniorchis to accommodate them. In 1996, Gustavo Romero and Germán Carnevali transferred to Bifrenaria a species originally described by Schlechter as Maxillaria petiolaris and now classified as Hylaeorchis petiolaris. On the same year, Vitorino Castro Neto published a revision of Bifrenaria, with five sections, which is the classification generally used today.
Bifrenaria has traditionally been classified in subtribe Bifrenariinae of tribe Maxillariae (Epidendroideae
), however, the relationships among the several genera within this tribe are not well defined and changes are expected in the upcoming years. The genus closest to Bifrenaria is Rudolfiella
. Other related genera are Teuscheria
, Guanchezia, Hylaeorchis and Horvatia
, in addition to the more distant Scuticaria
and Xylobium
. The unification of subtribes Lycastinae, Maxillariinae and Bifrenariinae has recently been suggested. However, there is no consensus on the path to be followed. Contrary to what was previously thought, the relationship among Bifrenaria and all these genera from Central America
seems to indicate a primitive origin of Bifrenaria in Central America
and its posterior dissemination towards the Southeast of Brazil, where it found fertile grounds to its more recent evolution.
In 2000, the first relatively complete molecular analysis on Bifrenaria species were made. Sixteen species from it and six of close genera were studied searching for confirmation of their phylogenetic relations, besides the delimitation of each species and each of Bifrenarias groups. The results did not allow for the acceptance of Adipe as a separate genus and, although they confirmed the monophyly of Cydoniorchis (B. tetragona and B. wittigii), they dissuade its recognition because six other genera would then be required to accommodate the remaining species. The study also expounded on convenience of splitting two species that are similar to each other and variable among themselves, but with many hard intermediate forms hard to delinetate as B. charlesworthii
and B. racemosa. It also confirmed the position of B. steyermarkii outside of Bifrenaria, but without suggesting a new name.
Bifrenaria is formed by about twenty species divided in two main groups of plants, large and small, with some visible morphological subdivisions highly confirmed by phylogeny.
Large species: is the group originally classified as Bifrenaria. They present four sided pseudobulb
s, with relatively short and erect inflorescence bearing up to ten fleshy large flowers but generally less. Usually the flowers are grouped and are fragrant or exhale strong scent. The labellum has three or four lobes and an elongated low callus. They are epiphytes, or often lithophytes. All originated in the southeast of Brazil. This group can be split in three subgroups:
Small species: is formed by the plants that once belonged to Stenocoryne, or more accurately, Adipe, which normally are epiphytes. They present smaller and not as noticeably four sided pseudobulbs, and long and delicate inflorescence bearing a higher medium number of flowers than the large species, although also never surpassing ten. The flowers are smaller and not fleshy, with an entire labellum, or sometimes slightly lobed on the apex. These species take less luminosoty and more humidity than those and are not particularly fragrant. According to their morphology they can be split in four distinct subgroups:
Other species: the remaining species are plants about which classification consensus has not been achieved: Bifrenaria maguirei, also classified under the genus Guanchezia, and Bifrenaria grandis, under Lacaena
. Bifrenaria steyermarkii
is a species highly different from all other Bifrenaria because its inflorescence is very long ant its flowers highly narrow, therefore it does not fit in any group, nevertheless the only other option of classification that has been published so far is under Xylobium
what possibly is not a choice either.
Bifrenaria exist from the north of South America, one species reaching Trinidad
, until Rio Grande do Sul
, the farther south State in Brazil
, however they are split in two isolated areas: Amazon Forest and Atlantic Forest of Brazil. The later, where seventeen species are present, may be considered their recente center of distribution. The montane area of Rio de Janeiro State and Espírito Santo
is particularly rich with fifteen species registered. Serra dos Órgãos
mountains area, in Rio, is reported as habitat of fourteen Bifrenaria species, however, some of these species are considered synonyms today, being eleven a more realistic number of species existing in the said area.
The species with large flowers are more common on Region Southeast of Brazil, however, they inhabit from the sunnier areas of the seashore to rocky mountain areas of Minas Gerais
e Bahia
States, from almost sea level up to 2,000 meters of altitude
, some species reaching Rio Grande do Sul
state. No species of large flowers exist in Amazon Forest. Some species grow directly attached to the famous Sugarloaf Mountain in Rio de Janeiro which can be observed by the commuters in the cable car. The recent centers of irradiation of this group are the seashore close to Serra do Mar
chain of mountains, and the high chains of mountains of Minas Gerais. The most common species in this group, spread from Rio Grande do Sul to Bahia, is B. harrisoniae
.
The smaller species of Bifrenaria, which some taxonomists classify under the genus Adipe, are more common on less sunny areas and can be found between 300 and about 1,600 meters of altitude. Six species are native in Serra do Mar Chain of mountains and its arms, place considered the center of distribution of the small species. Only three small species inhabit Amazon, B. longicornis
, which is more common at low artitudes; Bifrenaria venezuelana
, up to 1,450 meter of altitude, and B. steyermarkii
from even higher altitudes, in Roraima
State, in Brasil, and nearby areas in Venezuela
and Surinam.
The most common species is B. aureofulva
, however, because the geographic characteristic of its territory, without obstacles, B. longicornis
is the species spread throughout the largest area, reaching Colombia
, Venezuela
, Peru
, Surinam, Guyana
s, Trinidad and all Amazonic area in Brazil. Two species seem to be endemic in highly restricted areas: B. silvana
. discovered in 1987 at Serra da Ouricana mountains, nearby Itororó
, in Bahia
whch belongs to Adipe group; and B. verboonenii
, discovered in September 1995 on Serra do Cipó mountains, close to Diamantina, Minas Gerais
, of the large Bifrenara group.
Bifrenaria species inhabit three different environments. The large species generally live in well illuminated areas, occasionally epiphyte on trees of sparse foliage, more often as lithophyte
s, in campos rupestres, montane rocky areas that exist mostly in Rio and Minas Gerais States of Brazil, or over rocks in jungle's clearances. B. tyrianthina is exclusively lithophyte, B. tetragona and B. wittigii hardly ever do. B. atropurpurea is the only species found living terrestrially, but in rare occasions. The large species always show .
The small species from Southeast Brazil live in cloud montane forests, where the appear in much darker places than the large species. Within this sort of forests the temperature
presents noticeable difference between day and night and also through the seasons. These also are plants of caespitous growth, almost all epiphytes, despite there is at least one record of Bifrenaria aureofulva living lithophylically in Chapada Diamantina
, Bahia.
The species from Amazon inhabit tropical lowland
forests and equatorial forests. Bifrenaria longicornis is mostly found in flood areas along the igapós and igarapé
s (seasonal flood streams and small rivers of Amazon), and occasionally in open fields where the humidity
is high and temperature
constant through the year, normally in well illuminated places, although not under direct sunlight
. B. venezuelana inhabits forests in higher elevations, closer to the Andes
. Amazon species are epiphyte and the only Bifrenaria species with elongated rhyzome and ascendant growth.
Bifrenaria are comparatively easy to grow orchids. They should be preferably potted on well drained vegetable fiber because their roots and pseudobulbs rot easily when kept humid for long periods. One of three different environments is needed depending on the species' origin to successfully grow these plants. The larger species need more light than the others. The smaller species from Southeast Brazil may be cultivated at the same medium temperature but under less than 10-20% luminosity
. Bifrenaria from the Amazon Forest require higher and more constant temperature
and humidity
than other species. All species need most water
and fertilizer
during their active growth season.
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...
in family Orchidaceae
Orchidaceae
The Orchidaceae, commonly referred to as the orchid family, is a morphologically diverse and widespread family of monocots in the order Asparagales. Along with the Asteraceae, it is one of the two largest families of flowering plants, with between 21,950 and 26,049 currently accepted species,...
. It contains 20 species found in Panama
Panama
Panama , officially the Republic of Panama , is the southernmost country of Central America. Situated on the isthmus connecting North and South America, it is bordered by Costa Rica to the northwest, Colombia to the southeast, the Caribbean Sea to the north and the Pacific Ocean to the south. The...
, Trinidad
Trinidad
Trinidad is the larger and more populous of the two major islands and numerous landforms which make up the island nation of Trinidad and Tobago. It is the southernmost island in the Caribbean and lies just off the northeastern coast of Venezuela. With an area of it is also the fifth largest in...
and South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...
. There are no known uses for them, but their abundant, and at first glance artificial, 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, make them favorites of orchid growers.
The genus can be split in two clearly distinct groups: one of highly robust plants with large flowers, that encompass the first species to be classified under the genus Bifrenaria; other of more delicate plants with smaller flowers occasionally classified as Stenocoryne or Adipe. There are two additional species that are normally classified as Bifrenaria, but which molecular analysis indicate to belong to different orchid groups entirely. One is Bifrenaria grandis which is endemic to Bolívia
Bolivia
Bolivia officially known as Plurinational State of Bolivia , is a landlocked country in central South America. It is the poorest country in South America...
and which is now placed in Lacaena
Lacaena
Lacaena is a genus of orchids comprising six species and varieties native to Central America....
, and Bifrenaria steyermarkii
Bifrenaria steyermarkii
Bifrenaria steyermarkii is a species of orchid....
, an inhabitant of the northern Amazon Forest, which does not have an alternative classification.
Description
Sympodial
Sympodial means "with conjoined feet", and in biology is often used to refer to the outward morphology or mode of growth of organisms.-In botany:...
growth, between ten and sixty centimeters tall. They are characterized by round-section root with thick velamen
Velamen
Velamen is a spongy, multiple epidermis that covers the roots of some epiphytic or semi-epiphytic plants, such as orchid and Clivia species....
, four-angled fleshy pseudobulb
Pseudobulb
The pseudobulb is a storage organ derived from the part of a stem between two leaf nodes.It applies to the orchid family , specifically certain groups of epiphytic orchids, and may be single or composed of several internodes with evergreen or deciduous leaves along its length.In some species, it is...
s of one internode, often basally protected by dried sheaths and with only one apical leaf
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....
(except for Bifrenaria steyermarkii, which occasionally has two), plicate (fan-folded) enervated leathery leaves, yet malleable and not exceedingly thick, with a pseudo-petiole of basal round section, and a basal inflorescence
Inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...
s bearing up to ten flowers, which seldom surpass the leaves' length.
Bifrenaria flowers are strongly scented, they have sepals slightly larger than the petals, with the lateral ones basally united to the column
Column (botany)
The column, or technically the gynostemium, is a reproductive structure that can be found in several plant families: Aristolochiaceae, Orchidaceae, and Stylidiaceae....
foot forming a calcar
Calcar
The calcar, also known as the calcaneum , is the name given to a spur of cartilage arising from inner side of ankle and running along part of outer interfemoral membrane in bats , this is to help spread the interfemoral membrane , which is part of the wing membrane between the tail and the hind...
with truncated extremity.The column is slightly arching, generally without wings or any other appendages, bearing a foot which the labellum is hinged to, whose shape varies, articulated to the column, with a longitudinal channeled callus often with a basal claw. Flowers show two elongated stipe
Stipe (botany)
In botany, a stipe is a stalk that supports some other structure. The precise meaning is different depending on which taxonomic group is being described....
s, hardy ever one, at least twice longer than wide, with salient viscidium, visible caudicles and retinacle in inverted positions. The superposed pollinia number four, and are protected by a deciduous incumbent anther. Fruits are green, erect or pendulous; they take about eight months to ripe and hold hundreds of thousand yellowish or brownish elongated seeds up to 0.35 mm long. Among all the mentioned, the main characteristic distinguishing Bifrenaria from its closest relatives is the presence of the calcar. Other important characters are the four-sided single-leaved pseudobulbs besides the raceme inflorescence with two to ten flowers.
Little is known about pollination in Bifrenaria. Apparently the only existing records report the presence of some large species' pollinia observed on the back of male Eufriesea violacea bees (Euglossinae), and of Bombus brasiliensis (Bombini). Although there are no reports of flower pollination being directly observed, a paper published in 2006 studied the micromorphology of the labellum in Bifrenaria species, looking for substances useful to insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s as food. The absence of such substances on the densely pubescent surface of most Bifrenaria labelli seems to indicate possible pollination by large bees as the major mean. Another indicator of this possibility is the strong smell emanated by species like B. tetragona which are similar to those of plants in other families which are also pollinated by these bees. The smaller pubescent species may be pollinated by smaller bees, while the smooth ones, which have strong colored flowers, as B. aureofulva, might be pollinated by hummingbird
Hummingbird
Hummingbirds are birds that comprise the family Trochilidae. They are among the smallest of birds, most species measuring in the 7.5–13 cm range. Indeed, the smallest extant bird species is a hummingbird, the 5-cm Bee Hummingbird. They can hover in mid-air by rapidly flapping their wings...
s.
Taxonomy and phylogeny
The firstBifrenaria species to be described was in 1824 by English Botanist William Jackson HookerWilliam 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,...
, under the name Dendrobium harrisoniae
Bifrenaria harrisoniae
Bifrenaria harrisoniae is a species of orchid.-Synonyms:chronological:alphabetical:...
. Three years later, he also described the first small-flowered species, B. racemosa, but placed it in Maxillaria
Maxillaria
Maxillaria, abbreviated as Max in horticultural trade, is a large genus of orchids . This is a diverse genus, with very different morphological forms. Their characteristics can vary widely....
. With these two publications began a long series of species descriptions and confusing genera creation which generated much doubt for the next two centuries. The Royal Botanic Garden registers the submission of 69 species or subspecific taxa under Bifrenaria since the description of the first species. Among these, twenty are generally accepted but only seventeen are truly well established, with no doubts about their limits and classification. Thirteen other species are still accepted but now placed in other genera, and four or five, due to deficiencies in their descriptions, might never be definitely identified.
In 1832, John Lindley
John Lindley
John Lindley FRS was an English botanist, gardener and orchidologist.-Early years:Born in Catton, near Norwich, England, John Lindley was one of four children of George and Mary Lindley. George Lindley was a nurseryman and pomologist and ran a commercial nursery garden...
proposed the genus Bifrenaria and described its type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...
, Bifrenaria atropurpurea
Bifrenaria atropurpurea
Bifrenaria atropurpurea is a species of orchid and the type species of the genus Bifrenaria....
, previously named by Conrad Loddiges as Maxillaria atropurpurea. The name Bifrenaria comes from bi, two, and freno, brake, a reference to the shape of the two pairs of pollinia hold by separated caudicles presented by its flowers.
In 1837, Constantine Samuel Rafinesque, considering the noticeable vegetative difference between the few Bifrenaria known at the time, proposed the genus Adipe, based on B. racemosa morphology, described by Hooker few years before, to which he added the description of a supposed new species, Adipe fulva (today treated as a synonym of B. racemosa). The next year Lindley received a specimen from Amazonia, which was morphologically even more distant from the known species, but nonetheless described it as Bifrenaria longicornis
Bifrenaria longicornis
Bifrenaria longicornis is a species of orchid....
. Five years later, apparently not aware of Rafinesque's previous genus Adipe, Lindley changed his mind and suggested that this species should be classified under a new genus, Stenocoryne. Six species were subsequently attributed to Stenocoryne by various taxonomists, but the genus Rafinesque proposed remained unused until 1990.
Rudolfiella aurantiaca
Rudolfiella aurantiaca is a species of orchid occurring from Trinidad to tropical South America. It is the type species of the genus Rudolfiella....
(which presented the mentioned differences) as the type. However this genus name was already occupied (It is a synonym of Lindleya
Lindleya
Lindleya is a genus of Mexican evergreen trees of the family Rosaceae. The sole species, L. mespiloides, grows to a height of and bears solitary white fragrant flowers in summer....
, in the Rosaceae
Rosaceae
Rosaceae are a medium-sized family of flowering plants, including about 2830 species in 95 genera. The name is derived from the type genus Rosa. Among the largest genera are Alchemilla , Sorbus , Crataegus , Cotoneaster , and Rubus...
). Just thirty years later, in 1944, Frederico Carlos Hoehne, working on the first revision of genus Bifrenaria, corrected the suggestion of Schlechter. Hoehne initially proposed the genus Schlechterella for these species but, coincidentally, this name was also taken, this time by African Asclepiadaceae
Asclepiadaceae
According to APG II, the Asclepiadaceae is a former plant family now treated as a subfamily in the Apocynaceae...
. Finally a genus with an available name in homage to Schlechter was erected in the very next issue of the journal, Rudolfiella
Rudolfiella
Rudolfiella, abbreviated as Rud. in the horticultural trade, is a genus of orchids comprising eight species native to tropical South America and Panama....
, by which time time the number of species had increased to seven. On this revision, besides Rudolfiella, Hoehne divided Bifrenaria into two genera, accepting Lindley's Stenocoryne but calling attention to the existence of Rafinesque's Adipe, which should have nomenclatural priority, while also raising doubts about the identity of several described species. In 1990, Manfred Wolff formally resurrected the genus Adipe and transferred to it ten Bifrenaria species, besides the two already described by Rafinesque; his change was purely nomenclatural and he did not revisit the species.
Making the picture even more complex, in 1994, Karheinz Senghas, based on several characteristics shared only by B. tetragona and B. wittigii, described the genus Cydoniorchis to accommodate them. In 1996, Gustavo Romero and Germán Carnevali transferred to Bifrenaria a species originally described by Schlechter as Maxillaria petiolaris and now classified as Hylaeorchis petiolaris. On the same year, Vitorino Castro Neto published a revision of Bifrenaria, with five sections, which is the classification generally used today.
Bifrenaria has traditionally been classified in subtribe Bifrenariinae of tribe Maxillariae (Epidendroideae
Epidendroideae
In plant systematics Epidendroideae is a subfamily of the orchid family, Orchidaceae. Epidendroideae is larger than all the other orchid subfamilies together, comprising more than 15,000 species in 576 genera. Most Epidendroid orchids are tropical epiphytes, typically with pseudobulbs...
), however, the relationships among the several genera within this tribe are not well defined and changes are expected in the upcoming years. The genus closest to Bifrenaria is Rudolfiella
Rudolfiella
Rudolfiella, abbreviated as Rud. in the horticultural trade, is a genus of orchids comprising eight species native to tropical South America and Panama....
. Other related genera are Teuscheria
Teuscheria
Teuscheria is a genus of orchids comprising eight species native to Central America and South America. The genus is named for Henry Teuscher, an award-winning landscape artist and horticulturalist....
, Guanchezia, Hylaeorchis and Horvatia
Horvatia
Horvatia is a genus of flowering plants from the orchid family, Orchidaceae.- References :*Pridgeon, A.M., Cribb, P.J., Chase, M.A. & Rasmussen, F. eds. . Genera Orchidacearum 1. Oxford Univ. Press....
, in addition to the more distant Scuticaria
Scuticaria
Scuticaria is a genus of moray eel in the family Muraenidae.-Species:* Scuticaria okinawae * Scuticaria tigrina...
and Xylobium
Xylobium
Xylobium, abbreviated Xyl in horticultural trade, is a genus of plant in family Orchidaceae. It contains about 31 species native to tropical America....
. The unification of subtribes Lycastinae, Maxillariinae and Bifrenariinae has recently been suggested. However, there is no consensus on the path to be followed. Contrary to what was previously thought, the relationship among Bifrenaria and all these genera from Central America
Central America
Central America is the central geographic region of the Americas. It is the southernmost, isthmian portion of the North American continent, which connects with South America on the southeast. When considered part of the unified continental model, it is considered a subcontinent...
seems to indicate a primitive origin of Bifrenaria in Central America
Central America
Central America is the central geographic region of the Americas. It is the southernmost, isthmian portion of the North American continent, which connects with South America on the southeast. When considered part of the unified continental model, it is considered a subcontinent...
and its posterior dissemination towards the Southeast of Brazil, where it found fertile grounds to its more recent evolution.
In 2000, the first relatively complete molecular analysis on Bifrenaria species were made. Sixteen species from it and six of close genera were studied searching for confirmation of their phylogenetic relations, besides the delimitation of each species and each of Bifrenarias groups. The results did not allow for the acceptance of Adipe as a separate genus and, although they confirmed the monophyly of Cydoniorchis (B. tetragona and B. wittigii), they dissuade its recognition because six other genera would then be required to accommodate the remaining species. The study also expounded on convenience of splitting two species that are similar to each other and variable among themselves, but with many hard intermediate forms hard to delinetate as B. charlesworthii
Bifrenaria charlesworthii
Bifrenaria charlesworthii is a species of orchid....
and B. racemosa. It also confirmed the position of B. steyermarkii outside of Bifrenaria, but without suggesting a new name.
Species
Large species: is the group originally classified as Bifrenaria. They present four sided pseudobulb
Pseudobulb
The pseudobulb is a storage organ derived from the part of a stem between two leaf nodes.It applies to the orchid family , specifically certain groups of epiphytic orchids, and may be single or composed of several internodes with evergreen or deciduous leaves along its length.In some species, it is...
s, with relatively short and erect inflorescence bearing up to ten fleshy large flowers but generally less. Usually the flowers are grouped and are fragrant or exhale strong scent. The labellum has three or four lobes and an elongated low callus. They are epiphytes, or often lithophytes. All originated in the southeast of Brazil. This group can be split in three subgroups:
- The first subgroup is formed by plants that show a pollinarium with entire stipe and a salient callus just on its anterior region. The species are two, Bifrenaria calcarataBifrenaria calcarataBifrenaria calcarata is a species of orchid....
, That has the intermadiate lobe of the labellum approximately triangular and the lateral ones square, and B. mellicolorBifrenaria mellicolorBifrenaria mellicolor is a species of orchid....
, which has more rounded lobes. These species are similar and preferably epiphytes.
- The second subgroup also is formed by plants that show a pollinarium with entire stipe but the callus is entirely salient and fleshy. It is formed by the two species Senghas moved to genus Cydoniorchis, Bifrenaria tetragonaBifrenaria tetragonaBifrenaria tetragona is a species of orchid....
that presents a totally smooth labellum with rounded apex, and B. wittigiiBifrenaria wittigiiBifrenaria wittigii is a species of orchid....
, that has a partially pubescent labellum of acute apex.
- The third subgroup é formed by the four species with a bifurcated stipe: Bifrenaria atropurpureaBifrenaria atropurpureaBifrenaria atropurpurea is a species of orchid and the type species of the genus Bifrenaria....
, the only one with a cuneated viscidium; B. tyrianthinaBifrenaria tyrianthinaBifrenaria tyrianthina is a species of orchid....
, the only to have rounded viscidium; B. inodoraBifrenaria inodoraBifrenaria inodora is a species of orchid....
with greenish flowers and two lobed callus; and B. harrisoniaeBifrenaria harrisoniaeBifrenaria harrisoniae is a species of orchid.-Synonyms:chronological:alphabetical:...
, highly variable species of several colors, that always has three lobed callus on the labellum. The species of this group are often or exclusively lithophytes.
- Bifrenaria verbooneniiBifrenaria verbooneniiBifrenaria verboonenii is a species of orchid....
placement is not well defined so far, yet surely among the large species.
Small species: is formed by the plants that once belonged to Stenocoryne, or more accurately, Adipe, which normally are epiphytes. They present smaller and not as noticeably four sided pseudobulbs, and long and delicate inflorescence bearing a higher medium number of flowers than the large species, although also never surpassing ten. The flowers are smaller and not fleshy, with an entire labellum, or sometimes slightly lobed on the apex. These species take less luminosoty and more humidity than those and are not particularly fragrant. According to their morphology they can be split in four distinct subgroups:
- The first subgroup is formed by the two Amazonian species, with elongated rhyzome: Bifrenaria venezuelanaBifrenaria venezuelanaBifrenaria venezuelana is a species of orchid....
, that has short inflorescence and highly reduced calcar; and Bifrenaria longicornisBifrenaria longicornisBifrenaria longicornis is a species of orchid....
, with long inflorescence and noticeable calcar.
- Within the second subgroup there is only Bifrenaria aureofulvaBifrenaria aureofulvaBifrenaria aureofulva is a species of orchid....
, easily identifiable because of its bright orange flowers with acute sepals, petals and labellum, which do not open well.
- The third subgroup is formed by the two species which present petals que apresentam petals parallel to the column; These two species are hard to separate due to the high number of intermediate varieties: Bifrenaria charlesworthiiBifrenaria charlesworthiiBifrenaria charlesworthii is a species of orchid....
, that has more open and hairy flowers; and a Bifrenaria racemosaBifrenaria racemosaBifrenaria racemosa is a species of orchid....
.
- The last subgroup is formed by species that present petals oblique to the column. Two have petals and sepals marked with other colors: Bifrenaria clavigeraBifrenaria clavigeraBifrenaria clavigera is a species of orchid....
, that has a calcar formed by the fusion of the lateral sepals bases, and Bifrenaria silvanaBifrenaria silvanaBifrenaria silvana is a species of orchid....
where the calcar is a result of their superposition; Bifrenaria leucorhoda has white flowers with labellum veined of pink color. The two remaining species have highly similar yellow flowers, although of diverse sizes, the smaller and often paler is Bifrenaria stefanaeBifrenaria stefanaeBifrenaria stefanae is a species of orchid....
, the larger and of brighter color is Bifrenaria vitellinaBifrenaria vitellinaBifrenaria vitellina is a species of orchid....
.
Other species: the remaining species are plants about which classification consensus has not been achieved: Bifrenaria maguirei, also classified under the genus Guanchezia, and Bifrenaria grandis, under Lacaena
Lacaena
Lacaena is a genus of orchids comprising six species and varieties native to Central America....
. Bifrenaria steyermarkii
Bifrenaria steyermarkii
Bifrenaria steyermarkii is a species of orchid....
is a species highly different from all other Bifrenaria because its inflorescence is very long ant its flowers highly narrow, therefore it does not fit in any group, nevertheless the only other option of classification that has been published so far is under Xylobium
Xylobium
Xylobium, abbreviated Xyl in horticultural trade, is a genus of plant in family Orchidaceae. It contains about 31 species native to tropical America....
what possibly is not a choice either.
Distribution and habitat
Trinidad
Trinidad is the larger and more populous of the two major islands and numerous landforms which make up the island nation of Trinidad and Tobago. It is the southernmost island in the Caribbean and lies just off the northeastern coast of Venezuela. With an area of it is also the fifth largest in...
, until Rio Grande do Sul
Rio Grande do Sul
Rio Grande do Sul is the southernmost state in Brazil, and the state with the fifth highest Human Development Index in the country. In this state is located the southernmost city in the country, Chuí, on the border with Uruguay. In the region of Bento Gonçalves and Caxias do Sul, the largest wine...
, the farther south State in Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...
, however they are split in two isolated areas: Amazon Forest and Atlantic Forest of Brazil. The later, where seventeen species are present, may be considered their recente center of distribution. The montane area of Rio de Janeiro State and Espírito Santo
Espírito Santo
Espírito Santo is one of the states of southeastern Brazil, often referred to by the abbreviation "ES". Its capital is Vitória and the largest city is Vila Velha. The name of the state means literally "holy spirit" after the Holy Ghost of Christianity...
is particularly rich with fifteen species registered. Serra dos Órgãos
Serra dos Órgãos
Serra dos Órgãos is a mountain range in the state of Rio de Janeiro, Brazil, which was turned into a national park in 1939. It is located about a one-hour drive from the city of Rio de Janeiro....
mountains area, in Rio, is reported as habitat of fourteen Bifrenaria species, however, some of these species are considered synonyms today, being eleven a more realistic number of species existing in the said area.
The species with large flowers are more common on Region Southeast of Brazil, however, they inhabit from the sunnier areas of the seashore to rocky mountain areas of Minas Gerais
Minas Gerais
Minas Gerais is one of the 26 states of Brazil, of which it is the second most populous, the third richest, and the fourth largest in area. Minas Gerais is the Brazilian state with the largest number of Presidents of Brazil, the current one, Dilma Rousseff, being one of them. The capital is the...
e Bahia
Bahia
Bahia is one of the 26 states of Brazil, and is located in the northeastern part of the country on the Atlantic coast. It is the fourth most populous Brazilian state after São Paulo, Minas Gerais and Rio de Janeiro, and the fifth-largest in size...
States, from almost sea level up to 2,000 meters of altitude
Altitude
Altitude or height is defined based on the context in which it is used . As a general definition, altitude is a distance measurement, usually in the vertical or "up" direction, between a reference datum and a point or object. The reference datum also often varies according to the context...
, some species reaching Rio Grande do Sul
Rio Grande do Sul
Rio Grande do Sul is the southernmost state in Brazil, and the state with the fifth highest Human Development Index in the country. In this state is located the southernmost city in the country, Chuí, on the border with Uruguay. In the region of Bento Gonçalves and Caxias do Sul, the largest wine...
state. No species of large flowers exist in Amazon Forest. Some species grow directly attached to the famous Sugarloaf Mountain in Rio de Janeiro which can be observed by the commuters in the cable car. The recent centers of irradiation of this group are the seashore close to Serra do Mar
Serra do Mar
Serra do Mar is a 1,500 km long system of mountain ranges and escarpments in Southeastern Brazil, which runs in parallel to the Atlantic Ocean coast from the state of Espírito Santo to southern Santa Catarina, although some include Serra Geral in the Serra do Mar, in which case this range...
chain of mountains, and the high chains of mountains of Minas Gerais. The most common species in this group, spread from Rio Grande do Sul to Bahia, is B. harrisoniae
Bifrenaria harrisoniae
Bifrenaria harrisoniae is a species of orchid.-Synonyms:chronological:alphabetical:...
.
The smaller species of Bifrenaria, which some taxonomists classify under the genus Adipe, are more common on less sunny areas and can be found between 300 and about 1,600 meters of altitude. Six species are native in Serra do Mar Chain of mountains and its arms, place considered the center of distribution of the small species. Only three small species inhabit Amazon, B. longicornis
Bifrenaria longicornis
Bifrenaria longicornis is a species of orchid....
, which is more common at low artitudes; Bifrenaria venezuelana
Bifrenaria venezuelana
Bifrenaria venezuelana is a species of orchid....
, up to 1,450 meter of altitude, and B. steyermarkii
Bifrenaria steyermarkii
Bifrenaria steyermarkii is a species of orchid....
from even higher altitudes, in Roraima
Roraima
Roraima is the northernmost and least populated state of Brazil, located in the Amazon region. It borders the states of Amazonas and Pará, as well as the nations of Venezuela and Guyana. The population is 400,000 and the capital is Boa Vista...
State, in Brasil, and nearby areas in Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...
and Surinam.
The most common species is B. aureofulva
Bifrenaria aureofulva
Bifrenaria aureofulva is a species of orchid....
, however, because the geographic characteristic of its territory, without obstacles, B. longicornis
Bifrenaria longicornis
Bifrenaria longicornis is a species of orchid....
is the species spread throughout the largest area, reaching Colombia
Colombia
Colombia, officially the Republic of Colombia , is a unitary constitutional republic comprising thirty-two departments. The country is located in northwestern South America, bordered to the east by Venezuela and Brazil; to the south by Ecuador and Peru; to the north by the Caribbean Sea; to the...
, Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...
, 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....
, Surinam, Guyana
Guyana
Guyana , officially the Co-operative Republic of Guyana, previously the colony of British Guiana, is a sovereign state on the northern coast of South America that is culturally part of the Anglophone Caribbean. Guyana was a former colony of the Dutch and of the British...
s, Trinidad and all Amazonic area in Brazil. Two species seem to be endemic in highly restricted areas: B. silvana
Bifrenaria silvana
Bifrenaria silvana is a species of orchid....
. discovered in 1987 at Serra da Ouricana mountains, nearby Itororó
Itororó
Itororó is a town and municipality in the state of Bahia in the North-East region of Brazil.-References:...
, in Bahia
Bahia
Bahia is one of the 26 states of Brazil, and is located in the northeastern part of the country on the Atlantic coast. It is the fourth most populous Brazilian state after São Paulo, Minas Gerais and Rio de Janeiro, and the fifth-largest in size...
whch belongs to Adipe group; and B. verboonenii
Bifrenaria verboonenii
Bifrenaria verboonenii is a species of orchid....
, discovered in September 1995 on Serra do Cipó mountains, close to Diamantina, Minas Gerais
Minas Gerais
Minas Gerais is one of the 26 states of Brazil, of which it is the second most populous, the third richest, and the fourth largest in area. Minas Gerais is the Brazilian state with the largest number of Presidents of Brazil, the current one, Dilma Rousseff, being one of them. The capital is the...
, of the large Bifrenara group.
Lithophyte
Lithophytes are a type of plant that grows in or on rocks. Lithophytes feed off moss, nutrients in rain water, litter, and even their own dead tissue....
s, in campos rupestres, montane rocky areas that exist mostly in Rio and Minas Gerais States of Brazil, or over rocks in jungle's clearances. B. tyrianthina is exclusively lithophyte, B. tetragona and B. wittigii hardly ever do. B. atropurpurea is the only species found living terrestrially, but in rare occasions. The large species always show .
The small species from Southeast Brazil live in cloud montane forests, where the appear in much darker places than the large species. Within this sort of forests the temperature
Temperature
Temperature is a physical property of matter that quantitatively expresses the common notions of hot and cold. Objects of low temperature are cold, while various degrees of higher temperatures are referred to as warm or hot...
presents noticeable difference between day and night and also through the seasons. These also are plants of caespitous growth, almost all epiphytes, despite there is at least one record of Bifrenaria aureofulva living lithophylically in Chapada Diamantina
Chapada Diamantina
Chapada Diamantina is a region of Bahia state, in the Northeast of Brazil.The Chapada Diamantina has a dramatic landscape with high plains, table-top mesas, and steep cliffs or towers known as 'tepuy.' Before the arrival of the Portuguese in the 19th century, the only local inhabitants of the...
, Bahia.
The species from Amazon inhabit tropical lowland
Lowland
In physical geography, a lowland is any broad expanse of land with a general low level. The term is thus applied to the landward portion of the upward slope from oceanic depths to continental highlands, to a region of depression in the interior of a mountainous region, to a plain of denudation, or...
forests and equatorial forests. Bifrenaria longicornis is mostly found in flood areas along the igapós and igarapé
Igarapé
Igarapé is a Brazilian municipality located in the state of Minas Gerais. The city belongs to the mesoregion Metropolitana de Belo Horizonte and to the microregion of Belo Horizonte.-See also:* List of municipalities in Minas Gerais...
s (seasonal flood streams and small rivers of Amazon), and occasionally in open fields where the 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,...
is high and temperature
Temperature
Temperature is a physical property of matter that quantitatively expresses the common notions of hot and cold. Objects of low temperature are cold, while various degrees of higher temperatures are referred to as warm or hot...
constant through the year, normally in well illuminated places, although not under direct sunlight
Sunlight
Sunlight, in the broad sense, is the total frequency spectrum of electromagnetic radiation given off by the Sun. On Earth, sunlight is filtered through the Earth's atmosphere, and solar radiation is obvious as daylight when the Sun is above the horizon.When the direct solar radiation is not blocked...
. B. venezuelana inhabits forests in higher elevations, closer to the Andes
Andes
The Andes is the world's longest continental mountain range. It is a continual range of highlands along the western coast of South America. This range is about long, about to wide , and of an average height of about .Along its length, the Andes is split into several ranges, which are separated...
. Amazon species are epiphyte and the only Bifrenaria species with elongated rhyzome and ascendant growth.
Cultivation
Luminosity
Luminosity is a measurement of brightness.-In photometry and color imaging:In photometry, luminosity is sometimes incorrectly used to refer to luminance, which is the density of luminous intensity in a given direction. The SI unit for luminance is candela per square metre.The luminosity function...
. Bifrenaria from the Amazon Forest require higher and more constant temperature
Temperature
Temperature is a physical property of matter that quantitatively expresses the common notions of hot and cold. Objects of low temperature are cold, while various degrees of higher temperatures are referred to as warm or hot...
and 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,...
than other species. All species need most water
Water
Water is a chemical substance with the chemical formula H2O. A water molecule contains one oxygen and two hydrogen atoms connected by covalent bonds. Water is a liquid at ambient conditions, but it often co-exists on Earth with its solid state, ice, and gaseous state . Water also exists in a...
and fertilizer
Fertilizer
Fertilizer is any organic or inorganic material of natural or synthetic origin that is added to a soil to supply one or more plant nutrients essential to the growth of plants. A recent assessment found that about 40 to 60% of crop yields are attributable to commercial fertilizer use...
during their active growth season.