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'''''Leptotes''''' is a genus of [[orchid]]s formed by nine small species that grow in the dry jungles of south and southeast [[Brazil]], and also in [[Paraguay]] or [[Argentina]]. They are small [[epiphytic]] plants of [[caespitific]] growth, that sometimes resemble little ''[[Brassavola]]'', as they share the same type of thin terete leaves, however, they are more closely related to ''[[Loefgrenianthus]]''.
'''''Leptotes''''' is a genus of [[orchid]]s formed by nine small species that grow in the dry [[jungle]]s of south and southeast [[Brazil]], and also in [[Paraguay]] or [[Argentina]]. They are small [[epiphytic]] plants of [[caespitous growth]] that sometimes resemble little ''[[Brassavola]]'', as they share the same type of thin [[terete]] leaves, though they are more closely related to ''[[Loefgrenianthus]]''.


Some species of ''Leptotes'' are widely cultivated and form showy displays when completely in bloom although they are not among the easiest to grow. The majority of the species are not cultivated and some are so rare to be almost unknown; five of the species have only recently been described since 2000.  Beside being cultivated for their ornamental value, there are records of the flowers and fruits of ''[[Leptotes bicolor]]'' being used as a substitute for [[vanilla]] in milk, ice cream, tea and candies.<ref><span style="font-variant:small-caps;">Lawler, L.J.</span>: ''Ethnobotany of the Orchidaceae'' in '''Orchid biology: reviews and perspectives, Vol.3'''. J. Arditti Ed., Cornell University Press, Ithaca, New York, 1984.</ref>
Some species of ''Leptotes'' are widely cultivated and form showy displays when completely in bloom although they are not among the easiest to grow. The majority of the species are not cultivated and some are so rare to be almost unknown; five of the nine species have been described since 2000.  Besides being cultivated for their ornamental value, there are records of the flowers and fruits of ''[[Leptotes bicolor]]'' being used as a substitute for [[vanilla]] in [[milk]], [[ice cream]], [[tea]] and [[candy|candies]].<ref><span style="font-variant:small-caps;">Lawler, L.J.</span>: ''Ethnobotany of the Orchidaceae'' in '''Orchid biology: reviews and perspectives, Vol.3'''. J. Arditti Ed., Cornell University Press, Ithaca, New York, 1984.</ref>


==Distribution==
==Distribution==
{{Image|Sandstone cliffs close to Analandia.jpg|left|300px|Sandstone cliffs covered by some remaining patches of forest close to Analândia, São Paulo State, Brazil. It is the habitat of ''Leptotes pauloensis''.}}
{{Image|Sandstone cliffs close to Analandia.jpg|left|300px|Sandstone cliffs covered by some remaining patches of forest close to Analândia, São Paulo State, Brazil. This is the habitat of ''Leptotes pauloensis''.}}
The species of ''Leptotes'' were originally discovered in the [[Atlantic Forest]] of Brazil and are always seen living epiphytically. Two species have since been found in other countries, ''L. unicolor'' in Argentina, and ''L. bicolor'' in Paraguay. Three species show a high degree of [[endemism]] in the south of [[Bahia State]], these do not exist in any other place. The states of southeastern Brazil can be considered the center of its distribution since they host the highest number of species and the ''Leptotes'' are most abundant there, however, the range is from the Serra da Jibóia chain of mountains, in Bahia, to the North of [[Rio Grande do Sul State]].<ref name="CBGO"><span style="font-variant:small-caps;">Van den Berg, Cássio</span>: ''Leptotes'' in '''Genera Orchidacearum Vol.4''', pp. 271-3. Oxford University Press, 2006. ISBN 9780198507123.</ref> It is assumed that ''L. mogyensis'' is a variety collected by  Krakowizer found at Mogy das Cruzes, although this has not been verified as Christenson only worked with specimens under culture in the US.
The species of ''Leptotes'' were originally discovered in the [[Atlantic Forest]] of Brazil and are always seen living [[epiphyte|epiphytically]]. Two species have since been found in other countries, ''L. unicolor'' in Argentina, and ''L. bicolor'' in Paraguay. Three species show a high degree of [[endemism]] in the south of [[Bahia State]]. The states of southeastern Brazil can be considered the center of its distribution since they host the highest number of species and the ''Leptotes'' are most abundant there, however, the range is from the Serra da Jibóia chain of mountains, in Bahia, to the North of [[Rio Grande do Sul State]].<ref name="CBGO"><span style="font-variant:small-caps;">Van den Berg, Cássio</span>: ''Leptotes'' in '''Genera Orchidacearum Vol.4''', pp. 271-3. Oxford University Press, 2006. ISBN 9780198507123.</ref>  


The species in the group that are characterized by wide open flowers, such as ''Leptotes tenuis'' and ''L. pauloensis'', are more frequently found in cloud mountain forests. ''L. bicolor'' has the broadest range and can survive in both the cloud mountanin forests and the dryer woods on the plateaus of the [[Serra do Mar]] chain of mountains. ''Leptotes  unicolor'' grows optimally in cooler areas, above 700 meters of altitude, and is frequently found growing on ''[[Araucaria]]''  and ''[[Podocarpus]]'' trees in the southern areas of Brazil.<ref name="CBGO"/>
The species in the group that are characterized by wide open flowers, such as ''Leptotes tenuis'' and ''L. pauloensis'', are more frequently found in [[montane]] cloud forests. ''L. bicolor'' has the broadest range and can survive in both the cloud forests and the dryer woods on the plateaus of the [[Serra do Mar]] chain of mountains. ''Leptotes  unicolor'' grows optimally in cooler areas, above 700 meters of altitude, and is frequently found growing on ''[[Araucaria]]''  and ''[[Podocarpus]]'' trees in the southern areas of Brazil.<ref name="CBGO"/>


==Description==
==Description==
{{Image|Leptotes distribution map.png|left|200px|''Leptotes'' distribution map.}}
{{Image|Leptotes distribution map.png|left|200px|''Leptotes'' distribution map.}}
Species assigned to the genus ''Leptotes'' have a short [[rhizome]] of cylindrical section. They have small [[pseudobulb]]s that almost imperceptibly prolongate in one, rarely two, terete fleshy leaves. They have variable characteristics and can be short or long, erect or hanging, dark green or purple colors, frequently have a wrinkly surface and a deeper ridge in the face. The [[inflorescence]] is apical, generally short, and grows from the apex of the pseudobulb without a protecting [[spathe]] and bears up to seven flowers, although fewer are more common. The [[flower]]s are relatively large when compared to the overall plant dimensions, normally partially bent and in some species forming groups with a very showy aspect.<ref name="CBGO"/> They are fragrant and this perfume can last from ten to twenty days.<ref name="DM"><span style="font-variant:small-caps;">Miller, David; Richard Warren; Izabel Moura Miller & Helmut Seehawer</span>: '''Serra dos Órgãos sua história e suas orquídeas''', p. 240. Rio de Janeiro, 2006.</ref>  
Species assigned to the genus ''Leptotes'' have a short cylindrical [[rhizome]]. They have small [[pseudobulb]]s that almost imperceptibly prolongate in one, rarely two, terete fleshy leaves. They have variable characteristics and can be short or long, erect or hanging, dark green or purple, and often have a wrinkly surface and a deeper ridge in the face. The [[inflorescence]] is apical, generally short, and grows from the apex of the pseudobulb without a protecting [[spathe]] and bears up to seven flowers, although fewer are more common. The [[flower]]s are relatively large when compared to the overall plant dimensions, normally partially bent and in some species forming groups with a very showy aspect.<ref name="CBGO"/> They are fragrant and this perfume can last from ten to twenty days.<ref name="DM"><span style="font-variant:small-caps;">Miller, David; Richard Warren; Izabel Moura Miller & Helmut Seehawer</span>: '''Serra dos Órgãos sua história e suas orquídeas''', p. 240. Rio de Janeiro, 2006.</ref>  
{{Image|Leptotes.jpg|right|200px|Floral diagram for ''Leptotes''.}}
{{Image|Leptotes.jpg|right|200px|Floral diagram for ''Leptotes''.}}
The appearance of the [[petal]]s and [[sepal]]s is similar, both are elongated although the petals are slightly more narrow. Flower colors are generally greenish, white or variable shades of pink and the labellum petal can be spotted in pale yellow, light purple or lilac. The labellum is located along the [[Column (botanical)|column]] and trilobated (three lobes).  The lateral lobes are small and raised beside the column, although never involving it. The intermediate lobe is much bigger and quite variable between the species. They can have either lanceolate or obovate shape, occasionally be fleshy, flat or bending backwards; in some species they have denticulated edges but are smooth in others; rarely calli are present and it has claws which hold it attached to the sides of the column. The colomn is short, thick and erect, normally greenish, biauriculated, and bears six [[pollinia]] of different sizes, two larger in the center and four smaller in two pairs held by a short [[caudicle]] in an arrangement that is unique among orchids. Their [[fruit]]s are rounded, succulent and have a perfume reminiscent of  ''[[Vanilla (Orchid)|Vanilla]]''.<ref><span style="font-variant:small-caps;">Withner, Carl Leslie</span>: '''The Cattleyas and Their Relatives, Vol. 3''', pp.94-95. Timber Press, Oregon. ISBN 0881922692</ref>
The appearance of the [[petal]]s and [[sepal]]s is similar, both are elongated although the petals are slightly more narrow. Flower colors are generally greenish, white or variable shades of pink and the [[labellum]] (a special petal attractive to [[pollinator]]s) can be spotted in pale yellow, light purple or lilac. The labellum is located along the [[Column (botanical)|column]] and trilobed (three lobes).  The lateral lobes are small and raised beside the column, although never involving it. The intermediate lobe is much bigger and quite variable between species. They can have either lanceolate or obovate shape, occasionally be fleshy, flat or bending backwards; in some species they have denticulated edges but are smooth in others. Rarely, [[callus|calli]] are present, with claws holding them attached to the sides of the column. The column is short, thick and erect, normally greenish, biauriculated, and bears six [[pollinia]] of different sizes, two larger in the center and four smaller in two pairs held by a short [[caudicle]] in an arrangement that is unique among orchids. Their [[fruit]]s are rounded, succulent and have a perfume reminiscent of  ''[[Vanilla (Orchid)|Vanilla]]''.<ref><span style="font-variant:small-caps;">Withner, Carl Leslie</span>: '''The Cattleyas and Their Relatives, Vol. 3''', pp.94-95. Timber Press, Oregon. ISBN 0881922692</ref>


The agent for the [[pollination]] of ''Leptotes'' has never been observed. [[Cássio van den Berg]] postulates that bees are the primary agent due to the colors and morphology of its flowers<ref name="CBGO"/> while other orchidologists suspect pollination by [[hummingbird]]s is more important.<ref name="DM"/>
The agent for the [[pollination]] of ''Leptotes'' has never been observed. [[Cássio van den Berg]] postulates, judging from the colors and morphology of the flowers, that bees are the primary agent<ref name="CBGO"/>, while other orchidologists suspect pollination by [[hummingbird]]s is more important.<ref name="DM"/>
 
As the roots of ''Leptotes'' rot easily with excessive [[humidity]], the best results for their culture are achieved when they are mounted on plaques of vegetal fiber or tree cork. Watering and [[fertilizer]] must be more frequent during active growth periods and less during dormant periods. Ideal growth conditions require an intermediate temperature and exposure to filtered sunlight.<ref><span style="font-variant:small-caps;">Pridgeon, Alex</span>: ''Leptotes'' in '''Genera Orchidacearum Vol.4''', p. 274. Oxford Unity Press, 2006. ISBN 9780198507123.</ref>


==Taxonomic notes==  
==Taxonomic notes==  
{{Image|Leptotes bicolor Edwards.jpg|right|280px|Original illustration published by John Lindley when he described the genus ''Leptotes'', in 1833.}}
{{Image|Leptotes bicolor Edwards.jpg|right|280px|Original illustration published by John Lindley when he described the genus ''Leptotes'', in 1833.}}
In April of 1833, an unknown species from the [[Serra dos Órgãos]] mountains of Brazil blossomed in the greenhouse of Mrs. Arnold Harrison, an English collector of orchids homaged in diverse descriptions of noted species such as ''[[Bifrenaria harrisoniae]]'' and ''[[Cattleya harrisoniana]]''. A short time later, Mrs. Harrison sent a drawing and cutting of this plant to the botanist [[John Lindley]], who verified it to be not only a new species but a new genus. In its description, dated the same year, Lindley suggests the name of ''Leptotes'', from the [[Greek language |Greek]] for ''mild'', ''delicate'', in reference to the appearance of the plants flowers. He affirmed that ''Leptotes'' was similar to ''[[Tetramicra]]'', from which it is distinguished by the structures of the pollinarium and by the smaller lateral lobes of the lip petal; and because they have no calcar attached to the [[ovarium]]. He also differentiated it from  ''Brassavola'' by the pollinia and trilobed lip. Lindley then described its [[type-species]], ''[[Leptotes bicolor]]''.<ref><span style="font-variant:small-caps;">Lindley, John</span>: ''Leptotes bicolor'' in '''Edward's Botanical Register Vol.19''', t.1625. James Ridgway & Sons Ed. London, 1833. [http://www.botanicus.org/item/31753002748314 Published on internet.]</ref>   
In April of 1833, an unknown species from the [[Serra dos Órgãos]] mountains of Brazil blossomed in the greenhouse of Mrs. Arnold Harrison, an English collector of orchids homaged in diverse descriptions of noted species such as ''[[Bifrenaria harrisoniae]]'' and ''[[Cattleya harrisoniana]]''. A short time later, Mrs. Harrison sent a drawing and cutting of this plant to the botanist [[John Lindley]], who verified it to be not only a new species but a new genus. In its description, dated the same year, Lindley suggests the name of ''Leptotes'', from the [[Greek language |Greek]] λεπτότητα for ''mild'', ''delicate'', in reference to the appearance of the plant's flowers. He affirmed that ''Leptotes'' was similar to ''[[Tetramicra]]'', from which it is distinguished by the structures of the pollinarium and by the smaller lateral lobes of the lip petal; and because they have no calcar attached to the [[ovarium]]. He also differentiated it from  ''Brassavola'' by the pollinia and trilobed lip. Lindley then described its [[type-species]], ''[[Leptotes bicolor]]''.<ref><span style="font-variant:small-caps;">Lindley, John</span>: ''Leptotes bicolor'' in '''Edward's Botanical Register Vol.19''', t.1625. James Ridgway & Sons Ed. London, 1833. [http://www.botanicus.org/item/31753002748314 Published on internet.]</ref>   


In 1838, Lindley received two similar but slightly different plants, collected in [[Macaé de Cima]] and [[Ilha Grande]], in [[Rio de Janeiro]]. He considered them different to the previously described species, because the lateral lobes of the lip were slightly serrated, their flowers were bigger and they occasionally had a second leaf by the pseudobulb; he proposed a new species with the name of ''Leptotes serrulata''.<ref><span style="font-variant:small-caps;">Lindley, John</span>: ''Leptotes serrulata'' in '''Sertum orchidaceum''', t.11. James Ridgway & Sons Ed. London, 1838. [http://www.botanicus.org/item/31753002748314 Published on internet.]</ref> Five years later, the German Count [[Johann Centurius von Hoffmannsegg]] noticed that one plant he cultivated had a different green-bluish tone to its leaves and he described the plant calling it ''L. glaucophylla''.<ref><span style="font-variant:small-caps;">Hoffmannsegg, Johann Centurius von</span>: ''Leptotes glaucophylla'' in '''Botanische Zeitung  Vol.1''', p. 833. Berlin, 1843.</ref> When reviewing all the known species of ''Leptotes'' at the time, in 1990, [[Carl Leslie Withner]] reviewed the drawings published by Lindley and identified additional differences between ''L. bicolor'' and  ''L. serrulata'', describing how the latter always bears more flowers, with up to seven per inflorescence, which is also much longer.<ref name="W"><span style="font-variant:small-caps;">Withner, Carl Leslie</span>: '''The Cattleyas and Their Relatives, Vol. 3''', p.96. Timber Press, Oregon. ISBN 0881922692</ref> Today it is accepted, however, that both descriptions are variations of  ''L. bicolor'' that coexist due to its wide dispersion and multiplicity of populations.<ref name="K"><span style="font-variant:small-caps;">Govaerts, Rafaël ''et al''</span>: '''World Checklist of Orchidaceae'''. The Board of Trustees of the Royal Botanic Gardens, Kew. [http://www.kew.org/wcsp Published on Internet] (Access in January 2009).</ref>  
In 1838, Lindley received two similar but distinct plants, collected in [[Macaé de Cima]] and [[Ilha Grande]], in [[Rio de Janeiro]]. He considered them different to the previously described species, because the lateral lobes of the lip were slightly serrated, their flowers were bigger and they occasionally had a second leaf by the pseudobulb; he proposed a new species with the name of ''Leptotes serrulata''.<ref><span style="font-variant:small-caps;">Lindley, John</span>: ''Leptotes serrulata'' in '''Sertum orchidaceum''', t.11. James Ridgway & Sons Ed. London, 1838. [http://www.botanicus.org/item/31753002748314 Published on internet.]</ref> Five years later, the German Count [[Johann Centurius von Hoffmannsegg]] noticed that one plant he cultivated had a different green-bluish tone to its leaves and he described the plant calling it ''L. glaucophylla''.<ref><span style="font-variant:small-caps;">Hoffmannsegg, Johann Centurius von</span>: ''Leptotes glaucophylla'' in '''Botanische Zeitung  Vol.1''', p. 833. Berlin, 1843.</ref> When reviewing all the known species of ''Leptotes'' at the time, in 1990, [[Carl Leslie Withner|Carl Withner]] revisited the drawings published by Lindley and identified additional differences between ''L. bicolor'' and  ''L. serrulata'', describing how the latter always bears more flowers, with up to seven per inflorescence, which is also much longer.<ref name="W"><span style="font-variant:small-caps;">Withner, Carl Leslie</span>: '''The Cattleyas and Their Relatives, Vol. 3''', p.96. Timber Press, Oregon. ISBN 0881922692</ref> Today it is accepted, however, that both descriptions are variations of  ''L. bicolor'' that coexist due to its wide dispersion and multiplicity of populations.<ref name="K"><span style="font-variant:small-caps;">Govaerts, Rafaël ''et al''</span>: '''World Checklist of Orchidaceae'''. The Board of Trustees of the Royal Botanic Gardens, Kew. [http://www.kew.org/wcsp Published on Internet] (Access in January 2009).</ref>  


In 1865, [[Heinrich Gustav Reichenbach]], received, from some unknown locality in Brazil, the second described species to be accepted today, a plant much different from ''Leptotes bicolor'', because of its fewer and smaller rounded flowers, with wide open pale yellowish segments. He published it as ''L. tenuis''.<ref><span style="font-variant:small-caps;">Reichenbach, Heinrich Gustav</span>: ''Leptotes tenuis'' in '''Hamburger Garten Blumenzeitung Vol.21''', pp. 296. Hamburg, 1865.</ref> Twelve years later, the third species, ''Leptotes unicolor'', was described by Brazilian Botanist [[João Barbosa Rodrigues]]. According to the author, it was a much perfumed little species which he found some colonies with hundreds of plants living epiphyticaly along the banks of rivers [[Rio Dourado|Dourado]] and [[Rio Sapucai|Sapucai]], nearby the city of [[Alfenas]], in [[Minas Gerais]].<ref><span style="font-variant:small-caps;">Barbosa Rodrigues, João</span>: ''Leptotes unicolor'' in '''Genera et species orchidacearum novarum''' Vol.1, p.74. 1877. [http://www.botanicus.org/title/b12207019 Published on Internet, in French and Latin.]</ref> In 1881, Barbosa Rodrigues found another plant with longer leaves and slight diferences on floral structure, in [[Joinville]], a city that at the time belonged to [[Paraná]] State, but today is located in [[Santa Catarina]] which didn't exist then, and described it as ''L. paranaensis''.<ref><span style="font-variant:small-caps;">Barbosa Rodrigues, João</span>: ''Leptotes paranaensis'' in '''Genera et species orchidacearum novarum''' Vol.2, pp.163. 1881. [http://www.botanicus.org/title/b12207019 Published on Internet, in French and Latin.]</ref> Today this species is considered to be just a variation of ''L. unicolor'' he described four years earlier.<ref name="OB"><span style="font-variant:small-caps">Pabst, Guido & Dungs, Fritz </span>: '''Orchidaceae Brasilienses vol. 1''' p. 148, Brucke-Verlag Kurt Schmersow, Hildesheim, 1975. ISBN 3871050106 </ref>
In 1865, [[Heinrich Gustav Reichenbach]] received, from an unknown locality in Brazil, the second described species to be accepted today. The plant was quite different from ''Leptotes bicolor'', because of its fewer and smaller rounded flowers, with wide open pale yellowish segments and he published it as ''L. tenuis''.<ref><span style="font-variant:small-caps;">Reichenbach, Heinrich Gustav</span>: ''Leptotes tenuis'' in '''Hamburger Garten Blumenzeitung Vol.21''', pp. 296. Hamburg, 1865.</ref> Twelve years later, the third species, ''Leptotes unicolor'', was described by Brazilian botanist [[João Barbosa Rodrigues]]. His account described a highly fragrant little species of orchid and he found some colonies with hundreds of plants living epiphyticaly along the banks of the [[Rio Dourado|Dourado]] and [[Rio Sapucai|Sapucai]] rivers, nearby the city of [[Alfenas]], in [[Minas Gerais]].<ref><span style="font-variant:small-caps;">Barbosa Rodrigues, João</span>: ''Leptotes unicolor'' in '''Genera et species orchidacearum novarum''' Vol.1, p.74. 1877. [http://www.botanicus.org/title/b12207019 Published on Internet, in French and Latin.]</ref> In 1881, Barbosa Rodrigues found another species, this one with longer leaves and slight differences in floral structure, and named it as ''L. paranaensis'' after [[Paraná]] State where he first found the plant, although today the location near [[Joinville]] is located in [[Santa Catarina]].<ref><span style="font-variant:small-caps;">Barbosa Rodrigues, João</span>: ''Leptotes paranaensis'' in '''Genera et species orchidacearum novarum''' Vol.2, pp.163. 1881. [http://www.botanicus.org/title/b12207019 Published on Internet, in French and Latin.]</ref> Today this species is considered to be just a variation of ''L. unicolor'', the species he had described four years earlier.<ref name="OB"><span style="font-variant:small-caps">Pabst, Guido & Dungs, Fritz </span>: '''Orchidaceae Brasilienses vol. 1''' p. 148, Brucke-Verlag Kurt Schmersow, Hildesheim, 1975. ISBN 3871050106 </ref>


Eight years later, [[Robert Allen Rolfe]] received form Brazil, also without information of locality, a plant similar to ''Leptotes tenuis'' which Reichenbach described in 1865, however, with much thicker and shorter leaves. Rolfe described it as ''L. minuta''.<ref><span style="font-variant:small-caps;">Rolfe, Robert Allen</span>: ''Leptotes minuta'' in '''Gardeners' Chronicle  Vol.1889-2''':  p. 323. London, 1889.</ref> In his revision of Brazilian orchid species, published 1903, [[Célestin Alfred Cogniaux]], ignoring the variability of ''Leptotes'' species, both because of populations insolation and because of the variability of lenght and thickness of leaves due to diferent degrees of light exposure on each habitat, accepted this species which we know today is just a variation of ''L. tenuis''. Cogniaux did not have the opportunity to check the types of all other species then described, therefore, he accepted most of them with this remark.<ref><span style="font-variant:small-caps;">Cogniaux, Célestin Alfred</span>: ''Leptotes'' in '''Flora Brasiliensis Vol.3 Part.6''':  pp. 254-259. K.F.P. von Martius Ed., 1903. [http://florabrasiliensis.cria.org.br/search?taxon_id=1511 Published on Internet, in Latin.]</ref>
[[Robert Allen Rolfe]] received from Brazil, also without information of locality, a plant similar to the ''Leptotes tenuis'' species described by Reichenbach over 20 years earlier. Rolfe described it as ''L. minuta'' and noted it had much thicker and shorter leaves.<ref><span style="font-variant:small-caps;">Rolfe, Robert Allen</span>: ''Leptotes minuta'' in '''Gardeners' Chronicle  Vol.1889-2''':  p. 323. London, 1889.</ref> This new species was included in [[Célestin Alfred Cogniaux]]'s revision of Brazilian orchid species, published 1903, but in doing so he was unaware of the variability within the ''Leptotes'' species. At the time Cogniaux published his book he had not had the opportunity to check the types of all the other species then described, therefore, he accepted most of them with this remark.<ref><span style="font-variant:small-caps;">Cogniaux, Célestin Alfred</span>: ''Leptotes'' in '''Flora Brasiliensis Vol.3 Part.6''':  pp. 254-259. K.F.P. von Martius Ed., 1903. [http://florabrasiliensis.cria.org.br/search?taxon_id=1511 Published on Internet, in Latin.]</ref> In retrospect, it is now more clear that the variation in leaves was due to both the isolation of various populations and because of the different growth conditions in each habitat, and today ''L. minuta'' is known as a variation of ''L. tenuis''.


{{Image|Loefgrenianthus blancheamesiae.jpg|left|280px|''Loefgrenianthus blanche-amesiae'', first described under the genus ''Leptotes'', is its closest related species.}}
{{Image|Loefgrenianthus blancheamesiae.jpg|left|280px|''Loefgrenianthus blanche-amesiae'', first described under the genus ''Leptotes'', is its closest related species.}}
Johan Albert Constantin Loefgren, the Swedish Botanist living in Brazil, received from [[Itatiaia]], Rio de Janeiro, in 1918, one orchid with with pending habit, thin flat leaves, and flowers that rememinded the ones of ''Leptotes'', although its lips formed a pouch. He described it as ''Leptotes blanche-amesiae''.<ref><span style="font-variant:small-caps;">Loefgren, Johan Albert Constantin</span>: ''Leptotes blanche-amesiae'' in '''Arquivos do Jardim Botânico do Rio de Janeiro Vol.2''':  p. 58. Rio de Janeiro, 1918.</ref> Later [[Frederico Carlos Hoehne]] concluded this species, despite closely related, might be better placed in another genus thus, as an hommage to Loefgren, proposed the genus ''[[Loefgrenianthus]]'' to accomodate it.<ref><span style="font-variant:small-caps;">Hoehne, Frederico Carlos</span>: ''Loefgrenianthus'' in '''Arquivos de Botânica do Estado de São Paulo Vol.1''':  p. 593. São Paulo, Julho de 1927.</ref> Hoehne also described a new species, ''Leptotes pauloensis'', naming it so because is was found in [[São Paulo State]], in 1934. This species is close to  
While living in Brazil, the Danish Botanist [[Johan Albert Constantin Loefgren]] received an Orchid from [[Itatiaia]], Rio de Janeiro, with flowers reminiscent of the ''Leptotes'', although its lip petal formed a pouch. He described it as ''Leptotes blanche-amesiae'', also noting it had a pending habit and thin flat leaves.<ref><span style="font-variant:small-caps;">Loefgren, Johan Albert Constantin</span>: ''Leptotes blanche-amesiae'' in '''Arquivos do Jardim Botânico do Rio de Janeiro Vol.2''':  p. 58. Rio de Janeiro, 1918.</ref> Later work on the genus by [[Frederico Carlos Hoehne]] led him to conclude that this species, despite being closely related to the ''Leptotes'', would be better placed in another genus. He proposed the genus ''[[Loefgrenianthus]]'', in hommage to Loefgren.<ref><span style="font-variant:small-caps;">Hoehne, Frederico Carlos</span>: ''Loefgrenianthus'' in '''Arquivos de Botânica do Estado de São Paulo Vol.1''':  p. 593. São Paulo, Julho de 1927.</ref> In 1934, Hoehne also described a new species, ''Leptotes pauloensis'', naming it so because it was found in [[São Paulo State]]. This species is closely related to ''Leptotes tenuis'' but its flowers have different colors. For decades taxonomists were divided on whether this should be considered a new species, partly due to the fact it was so rare, indeed, [[Guido Frederico João Pabst|Guido Pabst]] considered it a synonym of ''L.  tenuis''.<ref name="OB"/> Recently many new colonies have been discovered and Withner now proposes that ''L. pauloensis'' should be accepted as a separate species.<ref name="W"/>
''Leptotes tenuis'' but has flowers of diferent colors. During decades taxonomists were not sure if this was to be considered a good species indeed. [[Guido Frederico João Pabst|Guido Pabst]] considered it as a synonym of the later;<ref name="OB"/> in 1990  Withner claimed that after being desappeared for many years, finally the doubts about its status were cleared as many had been found and it really should be accepted as good.<ref name="W"/>
 
Consequently, as of 2004, four species of ''Leptotes'' were known, three sufficiently different to be well established species, ''L. bicolor'', ''L. unicolor'' and ''L. tenuis'', and one, ''L. pauloensis'', that was becoming more frequently accepted by taxonomists.
 
A recent explosion of descriptions has more than doubled the size of this genus, however, the history of these discoveries starts much earlier. In 1954, one of the associates of [[Círculo Paulista de Orquidófilos]], an orchid society in São Paulo, presented a lecture where he talked informally about the innumerable varieties of ''Leptotes'' that he had collected throughout the years. This lecture was printed and distributed in the bulletin of the association.<ref><span style="font-variant:small-caps;">Krackowizer, F. J.</span>: ''Monografia do gênero Leptotes'' in '''Revista do Círculo Paulista de Orquidófilos Vol.11''', pp. 53-63 e 64-72. São Paulo, 1954.</ref> In 2004, [[Eric A. Christenson|Eric Christenson]] identified at least two of the several varieties mentioned in the lecture to be existing in collections throughout the [[United States of America]] and decided to describe them formally as independent species. One of these plants, ''Leptotes harryphillipsii'', is very similar to ''L. pauloensis'' already a problematic species by itself.<ref name="EC1"><span style="font-variant:small-caps;">Christenson, Eric A.</span>: ''Leptotes harryphillipsii'' in '''Orchids South Africa Vol.35''', pp. 53. Cape Town, 2004.</ref> The other one, ''L. mogyensis'', is unknown to Brazilian scholars and collectors. The sole example is the plant Christenson found in the US, supposedly originated from [[Mogi das Cruzes]], a city nearby São Paulo.<ref name="EC2"><span style="font-variant:small-caps;">Christenson, Eric A.</span>: ''Leptotes mogyensis'' in '''Orchids South Africa Vol.35''', pp. 54. Cape Town, 2004.</ref>


Therefore in 2004 four species of ''Leptotes'' were known, three sufficiently different to each other and well stablished, ''L. bicolor'', ''L. unicolor'' and ''L. tenuis'', and one fourth, ''L. pauloensis'', occasionally considered synonym of the previous one, but more frequently accepted by taxonomists. In this year, the species number doubled as other four were described. However, the history of these descriptions starts much earlier, in 1954, when one of the associates of [[Círculo Paulista de Orquidófilos]], an orchid society in São Paulo, presented a lecture where he talked long and informally about the innumerable varieties of ''Leptotes'' that he found and gathered throughout the years and the names he particularly atributed to each. This lecture was printed and distributed in the bulletin of the association.<ref><span style="font-variant:small-caps;">Krackowizer, F. J.</span>: ''Monografia do gênero Leptotes'' in '''Revista do Círculo Paulista de Orquidófilos Vol.11''', pp. 53-63 e 64-72. São Paulo, 1954.</ref> In 2004, [[Eric A. Christenson]], aware of this publication, found in culture in particular collections in the [[United States]] two of the several varieties mentioned in the lecture and decided to describe them formally as independent species. One of these plants, ''Leptotes harryphillipsii'', is very similar to ''L. pauloensis'' already a problematic species by itself.<ref name="EC1"><span style="font-variant:small-caps;">Christenson, Eric A.</span>: ''Leptotes harryphillipsii'' in '''Orchids South Africa Vol.35''', pp. 53. Cape Town, 2004.</ref> The other one, ''L. mogyensis'', is an unknown species to Brazilians, both scholar and collectors, except for the plant Christenson found in the US, supposedly originated from [[Mogi das Cruzes]], a city nearby São Paulo.<ref name="EC2"><span style="font-variant:small-caps;">Christenson, Eric A.</span>: ''Leptotes mogyensis'' in '''Orchids South Africa Vol.35''', pp. 54. Cape Town, 2004.</ref>
Two other new species described in 2004 belong to the affinity of ''Leptotes bicolor'' and were both found by the same surveyor in the region of [[Buerarema]], south of [[Bahia]]. ''Leptotes bohnkiana'', named after its finder, can be differentiated because it has a significantly smaller stature,<ref name="Campa"><span style="font-variant:small-caps;">Campacci, Marcos Antonio</span>: ''Leptotes bohnkiana'' in Boletim CAOB Vol.53, p.17. São Paulo, 2004. ISSN 14194590.</ref> the other, ''L. pohlitinocoi'', mostly by color.<ref><span style="font-variant:small-caps;">Vitorino P. Castro Neto & Chiron, Guy</span>: ''Leptotes pohlitinocoi'' in '''Richardiana Vol.4''': p.78. Paris, 2004.</ref>  


The other two species described in 2004 belong to the affinity of ''Leptotes bicolor'' and have been both found by the same surveyor in the region of [[Buerarema]], south of [[Bahia]]. ''Leptotes bohnkiana'' , which name is a homage to its finder, can be differentiated because of its smaller size,<ref name="Campa"><span style="font-variant:small-caps;">Campacci, Marcos Antonio</span>: ''Leptotes bohnkiana'' in Boletim CAOB Vol.53, p.17. São Paulo, 2004. ISSN 14194590.</ref> the other, ''L. pohlitinocoi'', mostly by color.<ref><span style="font-variant:small-caps;">Vitorino P. Castro Neto & Chiron, Guy</span>: ''Leptotes pohlitinocoi'' in '''Richardiana Vol.4''': p.78. Paris, 2004.</ref> Finally, in 2006 [[Sidney Marçal de Oliveira]] discovered the last species to be described, also from Bahia, although inhabitant of [[Chapada Diamantina]], ''Leptotes vellozicola'' is much diferent from all the other species.<ref name="CVB"><span style="font-variant:small-caps;">Van den Berg, Cássio ''et al''</span>: ''Leptotes vellozicola'' em '''Neodiversity Vol.1''', pp. 2, 2006. [http://www.neodiversity.org/articles/neod1101.pdf Published on internet.]</ref>
Finally, in 2006 [[Sidney Marçal de Oliveira]] discovered the last species to be described, also from Bahia, although an inhabitant of [[Chapada Diamantina]] too. This new species, ''Leptotes vellozicola'', is quite distinct from the other species.<ref name="CVB"><span style="font-variant:small-caps;">Van den Berg, Cássio ''et al''</span>: ''Leptotes vellozicola'' em '''Neodiversity Vol.1''', pp. 2, 2006. [http://www.neodiversity.org/articles/neod1101.pdf Published on internet.]</ref>


==Phylogeny==
According to [[Cássio van den Berg]] et al., who studied their phylogeny, ''Leptotes'' is very closely related to ''[[Loefgrenianthus]]'' and both situated between ''[[Pseudolaelia]]'' and the genus which once used to be classified as ''[[Schomburgkia]]'',  by some now considered part of ''[[Laelia]]''.<ref><span style="font-variant:small-caps;">Van den Berg, Cássio ''et al''</span>: ''A Phylogenetic analysis of Laellinae based on sequence data from internal transcribed spacers of nuclear ribosomal DNA'' in '''Lindleyana vol.15-2''', pp. 96–114, 2000. [http://www.cassiovandenberg.hpg.ig.com.br/pdfs/vandenbergetal2000.pdf Published on Internet].</ref>
According to [[Cássio van den Berg]] et al., ''Leptotes'' is very closely related to ''[[Loefgrenianthus]]'' and both situated between ''[[Pseudolaelia]]'' and the genus which once used to be classified as ''[[Schomburgkia]]'',  by some now considered part of ''[[Laelia]]''.<ref><span style="font-variant:small-caps;">Van den Berg, Cássio ''et al''</span>: ''A Phylogenetic analysis of Laellinae based on sequence data from internal trancribed spacers of nuclear ribosomal DNA'' in '''Lindleyana vol.15-2''', pp. 96–114, 2000. [http://www.cassiovandenberg.hpg.ig.com.br/pdfs/vandenbergetal2000.pdf Published on Internet].</ref>


==Species==
==Species==
{{Image|Leptotes bicolor blooming.jpg|right|280px|''Leptotes bicolor'' belongs to the group of species with long smooth leaves and bent elongated flowers that do not open well.}}The three main characteristics that differentiate between the species of ''Leptotes'' are the general proportions of the leaves, the shape of the flowers and way the flowers open. Using these differences the species can be classified into two main groups.
{{Image|Leptotes bicolor blooming.jpg|right|280px|''Leptotes bicolor'' belongs to the group of species with long smooth leaves and bent elongated flowers that do not open well.}}
The three main characteristics that differentiate between the species of ''Leptotes'' are the general proportions of the leaves, the shape of the flowers, and the way the flowers open. From these, the species can be classified into two main groups.


One group is formed by the four species with flowers of elongated segments, which generally are not widely open. These species use to present malleable inflorescences that leave the flowers slightly or very overthrown, frequent facing down. Almost all the species of this group have long leaves, of comparatively lighter tones, generally with smooth surfaces, longer than the inflorescences.
One group is formed by the four species with flowers of elongated segments, which generally are not widely open. These species present malleable inflorescences that leave the flowers slightly or very overthrown, frequently facing down. Almost all the species of this group have long leaves, of comparatively lighter tones, generally with smooth surfaces, that are longer than the inflorescences.


*''[[Leptotes unicolor]]'' is the exception in this group as it has short, wrinkly and dark leaves. Its flowers, of generally uniform pale pink, always face down. The other species of this group have flowers of stronger colors.<ref><span style="font-variant:small-caps;">Johnson, Andrés E.</span>: ''Leptotes unicolor'' in '''Las orquídeas del Parque Nacional Iguazú''', pp. 110. Fundación Vida Silvestre Argentina, Buenos Aires, 2001. ISBN 9509725412</ref>
*''[[Leptotes unicolor]]'' is the exception in this group, as it has short, wrinkly and dark leaves. Its flowers, of generally uniform pale pink, always face down. The other species of this group have flowers of stronger colors.<ref><span style="font-variant:small-caps;">Johnson, Andrés E.</span>: ''Leptotes unicolor'' in '''Las orquídeas del Parque Nacional Iguazú''', pp. 110. Fundación Vida Silvestre Argentina, Buenos Aires, 2001. ISBN 9509725412</ref>
*''[[Leptotes bicolor]]'' is the species with more flowers per inflorescence and with wider distribution. It is  a much variable species, even tho easily recognizeable because of its bicolored flowers, of white sepals and petals and purple lip, occasionally bearing two leaves per pseudobulb. There are some records of this species living as a lithophytic.<ref name="DM"/>
*''[[Leptotes bicolor]]'' is the species with more flowers per inflorescence and with a wider distribution. It is  a variable species, even though it is easily identified because of its bicolored flowers, white sepals and petals and purple lip; occasionally it will bear two leaves per pseudobulb. There are some records of this species living as a [[lithophyte]].<ref name="DM"/>
*''[[Leptotes bohnkiana]]'' is similar to ''L. bicolor'', however, its flowers measure only one third of the dimensions of the second ones, with segments proportionally wider, the plant measures only one half. All the  plants found had only one flower per inflorescence.  It just exists in Bahia.<ref name="Campa"/>
*''[[Leptotes bohnkiana]]'' has some similarities to ''L. bicolor'', however, its flowers are one third of the size, with petals and sepals that are proportionally wider, and the adult plant is about half the size. It bares a single flower per inflorescence and has only been found in Bahia.<ref name="Campa"/>
*''[[Leptotes pohlitinocoi]]'' is even closer to ''L. bicolor'' but has slightly smaller flowers with all segments completely pink. It just exists in Bahia.<ref><span style="font-variant:small-caps;">Vitorino P. Castro Neto</span>: ''Leptotes pohlitinocoi'' in '''Icones Orchidacearum Brasilienses Vol.2''': pl.163. São Paulo, March 2006. ISBN 8590149447</ref>
*''[[Leptotes pohlitinocoi]]'' is closer to ''L. bicolor'' but has slightly smaller flowers with all segments completely pink. It only exists in Bahia.<ref><span style="font-variant:small-caps;">Vitorino P. Castro Neto</span>: ''Leptotes pohlitinocoi'' in '''Icones Orchidacearum Brasilienses Vol.2''': pl.163. São Paulo, March 2006. ISBN 8590149447</ref>


The other group is formed by five smaller species with more rounded flowers of wide open flatter segments and much shorter wrinkly leaves, generally very dark green or purple colored. The species of this group often have only one or two flowers each inflorescence. Four of them are much similar and sometimes difficult to distinguish.
The other group is formed by five smaller species that have more rounded flowers with petals and sepals that are wide open and flatter. The leaves are shorter wrinkly leaves, generally very dark green or purple colored. The species of this group often have only one or two flowers on each inflorescence. Four of them are very similar and sometimes difficult to distinguish.
{{Image|Leptotes harryphillipsii plant.jpg|right|280px|''Leptotes harryphillipsii'' is a member of the ''Leptotes'' group of species with short wrinkled leaves and few wide open flowers.}}
{{Image|Leptotes harryphillipsii plant.jpg|right|280px|''Leptotes harryphillipsii'' is a member of the ''Leptotes'' group of species with short wrinkled leaves and few wide open flowers.}}
*''[[Leptotes vellozicola]]'' is the only easily recognizeable species in this group as it has a thick central callus next to the lip apex. It is the only species of this group endemic of Bahia, all the other are from Southeast and South Brazil.<ref name="CVB"/>
*''[[Leptotes vellozicola]]'' is the only easily recognizeable species in this group as it has a thick central callus next to the apex of the lip petal. It is the only species of this group endemic to Bahia, the other species are from southeast and south Brazil. This species, among all ''Leptotes'', takes the most sunlight in the wild because it is epiphytic on ''[[Vellozia]]'', a species with very few leaves. The region where it lives is much dryer compared to the areas inhabited by the other species.<ref name="CVB"/>
*''[[Leptotes tenuis]]'' is the only ''Leptotes'' species with pale green, yellowish or white flowers and lilac lip. It is a very small and uncommon species from Southeastern States of Brazil.<ref><span style="font-variant:small-caps;">Vitorino P. Castro Neto</span>: ''Leptotes tenuis'' in '''Icones Orchidacearum Brasilienses Vol.2''': pl.164. São Paulo, March 2006. ISBN 8590149447</ref>
*''[[Leptotes tenuis]]'' is the only ''Leptotes'' species with pale green, yellowish or white flowers and lilac colored lip. It is a very small and uncommon species from southeastern Brazil.<ref><span style="font-variant:small-caps;">Vitorino P. Castro Neto</span>: ''Leptotes tenuis'' in '''Icones Orchidacearum Brasilienses Vol.2''': pl.164. São Paulo, March 2006. ISBN 8590149447</ref>
*''[[Leptotes pauloensis]]'' is a species very similar to the previous one and can be separated from it mainly by the color of the floral segments, which are inverted here, that is, pale lilac petals and sepals and white lip with a yellowish cream mark in the middle. Its distribution overlaps ''L. tenuis'' but extends much more into South.<ref><span style="font-variant:small-caps;">Vitorino P. Castro Neto</span>: ''Leptotes pauloensis'' in '''Icones Orchidacearum Brasilienses Vol.2''': pl.162. São Paulo, March 2006. ISBN 8590149447</ref>
*''[[Leptotes pauloensis]]'' is a species very similar to ''L. tenuis'' and can be separated from it mainly by the opposite distribution of color between the lip and the other sepals and petals, namely, pale lilac petals and sepals and white lip with a yellowish cream mark in the middle. Its distribution overlaps ''L. tenuis'' but extends much more into the south.<ref><span style="font-variant:small-caps;">Vitorino P. Castro Neto</span>: ''Leptotes pauloensis'' in '''Icones Orchidacearum Brasilienses Vol.2''': pl.162. São Paulo, March 2006. ISBN 8590149447</ref>
*''[[Leptotes harryphillipsii]]'', another similar species to the previous, however with slightly longer lip and discrete pink stripes on the other segments, which shows a little more vibrant colors. It seems this species has been known for long time but has been always confused with the true ''L. pauloensis''.<ref name="EC1"/>
*''[[Leptotes harryphillipsii]]'', another species similar to ''L. tenuis'', but with a slightly longer lip and discrete pink stripes on the other petals and sepals, which typically have more vibrant colors. It seems this species has been known for long time but had always been confused with ''L. pauloensis''.<ref name="EC1"/>
*''[[Leptotes mogyensis]]'', yet another species related to ''L. tenuis'' group but that also somewhat resembles ''L. unicolor'' although bearing white flowers with a deep purple central mark on the lip. There is no record of this plant in the wild. All information comes from a plant found under cultivation in a nursery in California, USA. It might be a rare natural hibrid of the two mentioned species.<ref name="EC2"/>
*''[[Leptotes mogyensis]]'', yet another species related to the ''L. tenuis'' group and also resembles ''L. unicolor'' except it has white flowers with a deep purple central mark on the lip. There is no record of this plant in the wild. All information comes from a plant found under cultivation in a nursery in California, USA. It might be a rare natural hybrid of the two mentioned species.<ref name="EC2"/>
 
==Culture==
As the roots of ''Leptotes'' rot easily with excessive [[humidity]], the best results for their culture are achieved when they are mounted on plaques of vegetal fiber or tree cork. Watering and [[fertilizer]] must be more frequent during active growth periods and less during dormant periods. Ideal growth conditions require an intermediate temperature and exposure to filtered sunlight.<ref><span style="font-variant:small-caps;">Pridgeon, Alex</span>: ''Leptotes'' in '''Genera Orchidacearum Vol.4''', p. 274. Oxford Unity Press, 2006. ISBN 9780198507123.</ref>


==References==
==References==
{{reflist|2}}
{{reflist|2}}[[Category:Suggestion Bot Tag]]

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Leptotes
Leptotes bicolor
Leptotes bicolor
Scientific classification
Domain: Eukaryota
Kingdom: Plantae
Division: Magnoliophyta
Class: Liliopsida
Order: Asparagales
Family: Orchidaceae
Subfamily: Epidendroideae
Tribe: Epidendreae
Subtribe: Laeliinae
Genus: Leptotes
Lindl. 1833
Type species
Leptotes bicolor
Lindl. 1833
Species
Synonym

None

Leptotes is a genus of orchids formed by nine small species that grow in the dry jungles of south and southeast Brazil, and also in Paraguay or Argentina. They are small epiphytic plants of caespitous growth that sometimes resemble little Brassavola, as they share the same type of thin terete leaves, though they are more closely related to Loefgrenianthus.

Some species of Leptotes are widely cultivated and form showy displays when completely in bloom although they are not among the easiest to grow. The majority of the species are not cultivated and some are so rare to be almost unknown; five of the nine species have been described since 2000. Besides being cultivated for their ornamental value, there are records of the flowers and fruits of Leptotes bicolor being used as a substitute for vanilla in milk, ice cream, tea and candies.[1]

Distribution

(CC) Photo: Dalton Holland Baptista
Sandstone cliffs covered by some remaining patches of forest close to Analândia, São Paulo State, Brazil. This is the habitat of Leptotes pauloensis.

The species of Leptotes were originally discovered in the Atlantic Forest of Brazil and are always seen living epiphytically. Two species have since been found in other countries, L. unicolor in Argentina, and L. bicolor in Paraguay. Three species show a high degree of endemism in the south of Bahia State. The states of southeastern Brazil can be considered the center of its distribution since they host the highest number of species and the Leptotes are most abundant there, however, the range is from the Serra da Jibóia chain of mountains, in Bahia, to the North of Rio Grande do Sul State.[2]

The species in the group that are characterized by wide open flowers, such as Leptotes tenuis and L. pauloensis, are more frequently found in montane cloud forests. L. bicolor has the broadest range and can survive in both the cloud forests and the dryer woods on the plateaus of the Serra do Mar chain of mountains. Leptotes unicolor grows optimally in cooler areas, above 700 meters of altitude, and is frequently found growing on Araucaria and Podocarpus trees in the southern areas of Brazil.[2]

Description

(CC) Image: Dalton Holland Baptista
Leptotes distribution map.

Species assigned to the genus Leptotes have a short cylindrical rhizome. They have small pseudobulbs that almost imperceptibly prolongate in one, rarely two, terete fleshy leaves. They have variable characteristics and can be short or long, erect or hanging, dark green or purple, and often have a wrinkly surface and a deeper ridge in the face. The inflorescence is apical, generally short, and grows from the apex of the pseudobulb without a protecting spathe and bears up to seven flowers, although fewer are more common. The flowers are relatively large when compared to the overall plant dimensions, normally partially bent and in some species forming groups with a very showy aspect.[2] They are fragrant and this perfume can last from ten to twenty days.[3]

Floral diagram for Leptotes.

The appearance of the petals and sepals is similar, both are elongated although the petals are slightly more narrow. Flower colors are generally greenish, white or variable shades of pink and the labellum (a special petal attractive to pollinators) can be spotted in pale yellow, light purple or lilac. The labellum is located along the column and trilobed (three lobes). The lateral lobes are small and raised beside the column, although never involving it. The intermediate lobe is much bigger and quite variable between species. They can have either lanceolate or obovate shape, occasionally be fleshy, flat or bending backwards; in some species they have denticulated edges but are smooth in others. Rarely, calli are present, with claws holding them attached to the sides of the column. The column is short, thick and erect, normally greenish, biauriculated, and bears six pollinia of different sizes, two larger in the center and four smaller in two pairs held by a short caudicle in an arrangement that is unique among orchids. Their fruits are rounded, succulent and have a perfume reminiscent of Vanilla.[4]

The agent for the pollination of Leptotes has never been observed. Cássio van den Berg postulates, judging from the colors and morphology of the flowers, that bees are the primary agent[2], while other orchidologists suspect pollination by hummingbirds is more important.[3]

As the roots of Leptotes rot easily with excessive humidity, the best results for their culture are achieved when they are mounted on plaques of vegetal fiber or tree cork. Watering and fertilizer must be more frequent during active growth periods and less during dormant periods. Ideal growth conditions require an intermediate temperature and exposure to filtered sunlight.[5]

Taxonomic notes

Original illustration published by John Lindley when he described the genus Leptotes, in 1833.

In April of 1833, an unknown species from the Serra dos Órgãos mountains of Brazil blossomed in the greenhouse of Mrs. Arnold Harrison, an English collector of orchids homaged in diverse descriptions of noted species such as Bifrenaria harrisoniae and Cattleya harrisoniana. A short time later, Mrs. Harrison sent a drawing and cutting of this plant to the botanist John Lindley, who verified it to be not only a new species but a new genus. In its description, dated the same year, Lindley suggests the name of Leptotes, from the Greek λεπτότητα for mild, delicate, in reference to the appearance of the plant's flowers. He affirmed that Leptotes was similar to Tetramicra, from which it is distinguished by the structures of the pollinarium and by the smaller lateral lobes of the lip petal; and because they have no calcar attached to the ovarium. He also differentiated it from Brassavola by the pollinia and trilobed lip. Lindley then described its type-species, Leptotes bicolor.[6]

In 1838, Lindley received two similar but distinct plants, collected in Macaé de Cima and Ilha Grande, in Rio de Janeiro. He considered them different to the previously described species, because the lateral lobes of the lip were slightly serrated, their flowers were bigger and they occasionally had a second leaf by the pseudobulb; he proposed a new species with the name of Leptotes serrulata.[7] Five years later, the German Count Johann Centurius von Hoffmannsegg noticed that one plant he cultivated had a different green-bluish tone to its leaves and he described the plant calling it L. glaucophylla.[8] When reviewing all the known species of Leptotes at the time, in 1990, Carl Withner revisited the drawings published by Lindley and identified additional differences between L. bicolor and L. serrulata, describing how the latter always bears more flowers, with up to seven per inflorescence, which is also much longer.[9] Today it is accepted, however, that both descriptions are variations of L. bicolor that coexist due to its wide dispersion and multiplicity of populations.[10]

In 1865, Heinrich Gustav Reichenbach received, from an unknown locality in Brazil, the second described species to be accepted today. The plant was quite different from Leptotes bicolor, because of its fewer and smaller rounded flowers, with wide open pale yellowish segments and he published it as L. tenuis.[11] Twelve years later, the third species, Leptotes unicolor, was described by Brazilian botanist João Barbosa Rodrigues. His account described a highly fragrant little species of orchid and he found some colonies with hundreds of plants living epiphyticaly along the banks of the Dourado and Sapucai rivers, nearby the city of Alfenas, in Minas Gerais.[12] In 1881, Barbosa Rodrigues found another species, this one with longer leaves and slight differences in floral structure, and named it as L. paranaensis after Paraná State where he first found the plant, although today the location near Joinville is located in Santa Catarina.[13] Today this species is considered to be just a variation of L. unicolor, the species he had described four years earlier.[14]

Robert Allen Rolfe received from Brazil, also without information of locality, a plant similar to the Leptotes tenuis species described by Reichenbach over 20 years earlier. Rolfe described it as L. minuta and noted it had much thicker and shorter leaves.[15] This new species was included in Célestin Alfred Cogniaux's revision of Brazilian orchid species, published 1903, but in doing so he was unaware of the variability within the Leptotes species. At the time Cogniaux published his book he had not had the opportunity to check the types of all the other species then described, therefore, he accepted most of them with this remark.[16] In retrospect, it is now more clear that the variation in leaves was due to both the isolation of various populations and because of the different growth conditions in each habitat, and today L. minuta is known as a variation of L. tenuis.

(CC) Photo: Dalton Holland Baptista
Loefgrenianthus blanche-amesiae, first described under the genus Leptotes, is its closest related species.

While living in Brazil, the Danish Botanist Johan Albert Constantin Loefgren received an Orchid from Itatiaia, Rio de Janeiro, with flowers reminiscent of the Leptotes, although its lip petal formed a pouch. He described it as Leptotes blanche-amesiae, also noting it had a pending habit and thin flat leaves.[17] Later work on the genus by Frederico Carlos Hoehne led him to conclude that this species, despite being closely related to the Leptotes, would be better placed in another genus. He proposed the genus Loefgrenianthus, in hommage to Loefgren.[18] In 1934, Hoehne also described a new species, Leptotes pauloensis, naming it so because it was found in São Paulo State. This species is closely related to Leptotes tenuis but its flowers have different colors. For decades taxonomists were divided on whether this should be considered a new species, partly due to the fact it was so rare, indeed, Guido Pabst considered it a synonym of L. tenuis.[14] Recently many new colonies have been discovered and Withner now proposes that L. pauloensis should be accepted as a separate species.[9]

Consequently, as of 2004, four species of Leptotes were known, three sufficiently different to be well established species, L. bicolor, L. unicolor and L. tenuis, and one, L. pauloensis, that was becoming more frequently accepted by taxonomists.

A recent explosion of descriptions has more than doubled the size of this genus, however, the history of these discoveries starts much earlier. In 1954, one of the associates of Círculo Paulista de Orquidófilos, an orchid society in São Paulo, presented a lecture where he talked informally about the innumerable varieties of Leptotes that he had collected throughout the years. This lecture was printed and distributed in the bulletin of the association.[19] In 2004, Eric Christenson identified at least two of the several varieties mentioned in the lecture to be existing in collections throughout the United States of America and decided to describe them formally as independent species. One of these plants, Leptotes harryphillipsii, is very similar to L. pauloensis already a problematic species by itself.[20] The other one, L. mogyensis, is unknown to Brazilian scholars and collectors. The sole example is the plant Christenson found in the US, supposedly originated from Mogi das Cruzes, a city nearby São Paulo.[21]

Two other new species described in 2004 belong to the affinity of Leptotes bicolor and were both found by the same surveyor in the region of Buerarema, south of Bahia. Leptotes bohnkiana, named after its finder, can be differentiated because it has a significantly smaller stature,[22] the other, L. pohlitinocoi, mostly by color.[23]

Finally, in 2006 Sidney Marçal de Oliveira discovered the last species to be described, also from Bahia, although an inhabitant of Chapada Diamantina too. This new species, Leptotes vellozicola, is quite distinct from the other species.[24]

According to Cássio van den Berg et al., who studied their phylogeny, Leptotes is very closely related to Loefgrenianthus and both situated between Pseudolaelia and the genus which once used to be classified as Schomburgkia, by some now considered part of Laelia.[25]

Species

(CC) Photo: Dalton Holland Baptista
Leptotes bicolor belongs to the group of species with long smooth leaves and bent elongated flowers that do not open well.

The three main characteristics that differentiate between the species of Leptotes are the general proportions of the leaves, the shape of the flowers, and the way the flowers open. From these, the species can be classified into two main groups.

One group is formed by the four species with flowers of elongated segments, which generally are not widely open. These species present malleable inflorescences that leave the flowers slightly or very overthrown, frequently facing down. Almost all the species of this group have long leaves, of comparatively lighter tones, generally with smooth surfaces, that are longer than the inflorescences.

  • Leptotes unicolor is the exception in this group, as it has short, wrinkly and dark leaves. Its flowers, of generally uniform pale pink, always face down. The other species of this group have flowers of stronger colors.[26]
  • Leptotes bicolor is the species with more flowers per inflorescence and with a wider distribution. It is a variable species, even though it is easily identified because of its bicolored flowers, white sepals and petals and purple lip; occasionally it will bear two leaves per pseudobulb. There are some records of this species living as a lithophyte.[3]
  • Leptotes bohnkiana has some similarities to L. bicolor, however, its flowers are one third of the size, with petals and sepals that are proportionally wider, and the adult plant is about half the size. It bares a single flower per inflorescence and has only been found in Bahia.[22]
  • Leptotes pohlitinocoi is closer to L. bicolor but has slightly smaller flowers with all segments completely pink. It only exists in Bahia.[27]

The other group is formed by five smaller species that have more rounded flowers with petals and sepals that are wide open and flatter. The leaves are shorter wrinkly leaves, generally very dark green or purple colored. The species of this group often have only one or two flowers on each inflorescence. Four of them are very similar and sometimes difficult to distinguish.

(CC) Photo: Dalton Holland Baptista
Leptotes harryphillipsii is a member of the Leptotes group of species with short wrinkled leaves and few wide open flowers.
  • Leptotes vellozicola is the only easily recognizeable species in this group as it has a thick central callus next to the apex of the lip petal. It is the only species of this group endemic to Bahia, the other species are from southeast and south Brazil. This species, among all Leptotes, takes the most sunlight in the wild because it is epiphytic on Vellozia, a species with very few leaves. The region where it lives is much dryer compared to the areas inhabited by the other species.[24]
  • Leptotes tenuis is the only Leptotes species with pale green, yellowish or white flowers and lilac colored lip. It is a very small and uncommon species from southeastern Brazil.[28]
  • Leptotes pauloensis is a species very similar to L. tenuis and can be separated from it mainly by the opposite distribution of color between the lip and the other sepals and petals, namely, pale lilac petals and sepals and white lip with a yellowish cream mark in the middle. Its distribution overlaps L. tenuis but extends much more into the south.[29]
  • Leptotes harryphillipsii, another species similar to L. tenuis, but with a slightly longer lip and discrete pink stripes on the other petals and sepals, which typically have more vibrant colors. It seems this species has been known for long time but had always been confused with L. pauloensis.[20]
  • Leptotes mogyensis, yet another species related to the L. tenuis group and also resembles L. unicolor except it has white flowers with a deep purple central mark on the lip. There is no record of this plant in the wild. All information comes from a plant found under cultivation in a nursery in California, USA. It might be a rare natural hybrid of the two mentioned species.[21]

References

  1. ↑ Lawler, L.J.: Ethnobotany of the Orchidaceae in Orchid biology: reviews and perspectives, Vol.3. J. Arditti Ed., Cornell University Press, Ithaca, New York, 1984.
  2. ↑ 2.0 2.1 2.2 2.3 Van den Berg, Cássio: Leptotes in Genera Orchidacearum Vol.4, pp. 271-3. Oxford University Press, 2006. ISBN 9780198507123.
  3. ↑ 3.0 3.1 3.2 Miller, David; Richard Warren; Izabel Moura Miller & Helmut Seehawer: Serra dos Órgãos sua história e suas orquídeas, p. 240. Rio de Janeiro, 2006.
  4. ↑ Withner, Carl Leslie: The Cattleyas and Their Relatives, Vol. 3, pp.94-95. Timber Press, Oregon. ISBN 0881922692
  5. ↑ Pridgeon, Alex: Leptotes in Genera Orchidacearum Vol.4, p. 274. Oxford Unity Press, 2006. ISBN 9780198507123.
  6. ↑ Lindley, John: Leptotes bicolor in Edward's Botanical Register Vol.19, t.1625. James Ridgway & Sons Ed. London, 1833. Published on internet.
  7. ↑ Lindley, John: Leptotes serrulata in Sertum orchidaceum, t.11. James Ridgway & Sons Ed. London, 1838. Published on internet.
  8. ↑ Hoffmannsegg, Johann Centurius von: Leptotes glaucophylla in Botanische Zeitung Vol.1, p. 833. Berlin, 1843.
  9. ↑ 9.0 9.1 Withner, Carl Leslie: The Cattleyas and Their Relatives, Vol. 3, p.96. Timber Press, Oregon. ISBN 0881922692
  10. ↑ Govaerts, Rafaël et al: World Checklist of Orchidaceae. The Board of Trustees of the Royal Botanic Gardens, Kew. Published on Internet (Access in January 2009).
  11. ↑ Reichenbach, Heinrich Gustav: Leptotes tenuis in Hamburger Garten Blumenzeitung Vol.21, pp. 296. Hamburg, 1865.
  12. ↑ Barbosa Rodrigues, João: Leptotes unicolor in Genera et species orchidacearum novarum Vol.1, p.74. 1877. Published on Internet, in French and Latin.
  13. ↑ Barbosa Rodrigues, João: Leptotes paranaensis in Genera et species orchidacearum novarum Vol.2, pp.163. 1881. Published on Internet, in French and Latin.
  14. ↑ 14.0 14.1 Pabst, Guido & Dungs, Fritz : Orchidaceae Brasilienses vol. 1 p. 148, Brucke-Verlag Kurt Schmersow, Hildesheim, 1975. ISBN 3871050106
  15. ↑ Rolfe, Robert Allen: Leptotes minuta in Gardeners' Chronicle Vol.1889-2: p. 323. London, 1889.
  16. ↑ Cogniaux, Célestin Alfred: Leptotes in Flora Brasiliensis Vol.3 Part.6: pp. 254-259. K.F.P. von Martius Ed., 1903. Published on Internet, in Latin.
  17. ↑ Loefgren, Johan Albert Constantin: Leptotes blanche-amesiae in Arquivos do Jardim Botânico do Rio de Janeiro Vol.2: p. 58. Rio de Janeiro, 1918.
  18. ↑ Hoehne, Frederico Carlos: Loefgrenianthus in Arquivos de Botânica do Estado de São Paulo Vol.1: p. 593. São Paulo, Julho de 1927.
  19. ↑ Krackowizer, F. J.: Monografia do gênero Leptotes in Revista do Círculo Paulista de Orquidófilos Vol.11, pp. 53-63 e 64-72. São Paulo, 1954.
  20. ↑ 20.0 20.1 Christenson, Eric A.: Leptotes harryphillipsii in Orchids South Africa Vol.35, pp. 53. Cape Town, 2004.
  21. ↑ 21.0 21.1 Christenson, Eric A.: Leptotes mogyensis in Orchids South Africa Vol.35, pp. 54. Cape Town, 2004.
  22. ↑ 22.0 22.1 Campacci, Marcos Antonio: Leptotes bohnkiana in Boletim CAOB Vol.53, p.17. São Paulo, 2004. ISSN 14194590.
  23. ↑ Vitorino P. Castro Neto & Chiron, Guy: Leptotes pohlitinocoi in Richardiana Vol.4: p.78. Paris, 2004.
  24. ↑ 24.0 24.1 Van den Berg, Cássio et al: Leptotes vellozicola em Neodiversity Vol.1, pp. 2, 2006. Published on internet.
  25. ↑ Van den Berg, Cássio et al: A Phylogenetic analysis of Laellinae based on sequence data from internal transcribed spacers of nuclear ribosomal DNA in Lindleyana vol.15-2, pp. 96–114, 2000. Published on Internet.
  26. ↑ Johnson, Andrés E.: Leptotes unicolor in Las orquídeas del Parque Nacional Iguazú, pp. 110. Fundación Vida Silvestre Argentina, Buenos Aires, 2001. ISBN 9509725412
  27. ↑ Vitorino P. Castro Neto: Leptotes pohlitinocoi in Icones Orchidacearum Brasilienses Vol.2: pl.163. São Paulo, March 2006. ISBN 8590149447
  28. ↑ Vitorino P. Castro Neto: Leptotes tenuis in Icones Orchidacearum Brasilienses Vol.2: pl.164. São Paulo, March 2006. ISBN 8590149447
  29. ↑ Vitorino P. Castro Neto: Leptotes pauloensis in Icones Orchidacearum Brasilienses Vol.2: pl.162. São Paulo, March 2006. ISBN 8590149447
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