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| <!-- supporters            --> [[User:Caesar Schinas|Caesar Schinas]] 09:25, 8 June 2009 (UTC);<BR/> [[User:Meg Ireland|Meg Ireland]] 08:00, 18 June 2009 (UTC)
| <!-- supporters            --> [[User:Caesar Schinas|Caesar Schinas]] 09:25, 8 June 2009 (UTC);<BR/> [[User:Meg Ireland|Meg Ireland]] 08:00, 18 June 2009 (UTC)

Revision as of 19:30, 24 June 2009

The Article of the Week is an article chosen by vote among Citizens as exemplifying various qualities we like to see in a Citizendium article; see our article standards.

Add New Nominees Here

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Table of Nominees
Nominated article Vote
Score
Supporters Specialist supporters
Developed Article Scuticaria 4 Caesar Schinas 09:25, 8 June 2009 (UTC);
Meg Ireland 08:00, 18 June 2009 (UTC)
Daniel Mietchen 17:24, 4 June 2009 (UTC)
Torture 5 Milton Beychok 01:33, 19 June 2009 (UTC);
Drew R. Smith 03:58, 19 June 2009 (UTC);
Peter Schmitt 23:23, 24 June 2009 (UTC)
Howard C. Berkowitz 14:32, 5 June 2009 (UTC)
Developed Article Milpa agriculture 3 Milton Beychok 00:15, 29 May 2009 (UTC);
Arne Eickenberg 14:15, 17 June 2009 (UTC);
Peter Schmitt 23:23, 24 June 2009 (UTC)

Transclusion of the above nominees (to be done by an Administrator)

View Current Transcluded Nominees (after they have been transcluded by an Administrator)

The next article (or draft) of the week will be the article with the most votes at 1 AM UTC on Thursday, 25 June 2009. I did the honors this time. Milton Beychok 06:18, 18 June 2009 (UTC)

Text in this section is transcluded from the respective Citizendium entries and may change when these are edited.

Nominated article Supporters Specialist supporters Score
Developed Article Miltonia: An orchid genus formed by nine showy epiphyte species and seven natural hybrids of Brazil, one species reaching Argentina and Paraguay. [e]
Miltonia
Miltonia spectabilis
Miltonia spectabilis
Scientific classification
Domain: Eukaryota
Kingdom: Plantae
Division: Magnoliophyta
Class: Liliopsida
Order: Asparagales
Family: Orchidaceae
Subfamily: Epidendroideae
Tribe: Cymbidieae
Subtribe: Oncidiinae
Genus: Miltonia
Lindl. 1837
Type species
Miltonia spectabilis
Lindl. 1837
Species
Synonyms

Miltonia is an orchid genus formed by nine epiphyte species and eight natural hybrids inhabitants of Brazilian Atlantic Forest, one species reaching the northeast of Argentina and east of Paraguay. This genus was established by John Lindley in 1837, when he described its type species, Miltonia spectabilis. Many species were attributed to Miltonia in the past, however, today, the species from Central America and from cooler areas on northwest of South America have been moved to other genera. Miltonia species have large and long lasting flowers, often in inflorescences with several of them. This fact, allied to being species that are easy to grow and to identify, make them a favorite of orchid collectors all over the world. Species of this genus are extensively used to produce artificial hybrids.

Despite the fact that Miltonia is now a well established genus, most of its species were originally classified under other genera as Cyrtochilum, Oncidium, Odondoglossum, and Brassia. All were discovered between 1834 and 1850 with the exception of M. kayasimae, discovered only in 1976.

Distribution

(CC) Photo: Dalton Holland Baptista
Miltonia candida
This is the only species with a labellum that embraces the column in a way that reminds Cattleya species.

Miltonia species range starts on the area of Missiones in the northeast of Argentina[1] and east of Paraguay[2] and spreads north along the Brazilian mountains of Serra do Mar and its branches up to the State of Pernambuco on Brazilian northeast. They occupy mostly areas between 200 and 1,500 meters of altitude meters, however the majority of the species are more often found about 600 to 900 meters. Miltonia species can be found from shady areas inside the forest to areas more exposed to the sun, however never are under full sunlight; usually in ventilated places where they receive plenty humidity during the night and early morning. They are always epiphyte and, because they grow very fast, each pseudobulb originating two new growths every year, they soon form large colonies.[3]

Miltonia russelliana and M. flavescens are the ones with the widest dispersion and found at lower altitudes. M. flavescens is the only species that exists in countries other than Brazil and is also the one that spreads farther north. M russelliana range starts on Rio Grande do Sul and ends at Bahia State. M. regnellii is also widespread although does not go northern than Rio de Janeiro. M. moreliana is a species more common at lower altitudes and warmer areas existing from Rio to Pernambuco. Miltonia candida, M. clowesii and M. spectabilis are restricted to the four states of Region Southeast of Brazil. Miltonia cuneata is just from São Paulo and Rio and the one that grow at highest altitudes. M. kayasimae is the only species really rare; it has been found just a couple of times in a very restricted area close to Salesópolis, in São Paulo State. The mountains area between São Paulo and Rio de Janeiro, where almost all species do exist may be considered the center of distribution of Miltonia.[4]

Description

(CC) Photo: Dalton Holland Baptista
Miltonia regnellii
This species shows the widest flower color variation among Miltonia species; they can vary from white to yellow, pink and lilac with labelli also from white to dark purple.

Miltonia are comparatively medium large orchid plants reaching about fifty centimeters height. They present subcaespitous growth, that means their pseudobulbs are not tightly packed but slightly spaced by a rhyzome, that is longer than on caespitous plants, with length between two and five centimeters. Their roots grow along the rhyzome in high numbers. They are white, comparatively thin, usually short and hardly branched. The rhyzome is covered by dried imbricating steaths which get increasingly larger at the base of pseudobulb becoming articulated foliar steaths that partially cover them. The pseudobulbs and leaves vary in color from yellowish bright light green to olive green depending on the species and to the amount of sunlight they are exposed to. They may be more oval and laterally highly flattened to slightly tetragonal and elongated and almost always bear two apical leaves. The leaves are narrow, flexible and hardly larger than three centimeters wide and forty long with the apexes rounded sometimes slightly pointed. Some species are about half of this size. The inflorescences are one or two per pseudobulb, shoot from their bases behind the protecting steaths. They are erect and never branched, often longer than the leaves, bearing from one to twelve moderately spaced flowers that open at the same time or in succession holding three or four opened all the time, when the older fades a new one opens. The older flowers of species with white lips that open in succession usually get yellower about the time the next flower opens although they still last one more week before fading. The first to bloom is M. cuneata, during late winter, but the majority of species bloom from late spring to late summer.[5]

The flowers of Miltonia vary from four to fifteen centimeters across; the larger are the ones with fewer flowers. Their colors vary from entirely white and pink to dark purple, pale yellow or lilac when plain, or they may highly spotted but then usually they are greenish or brownish with a contrasting labellum often white with purple dots, stains or veins close to the base.[3] The petals and sepals shapes are highly variable from species to species but always somewhat similar to each other within a species. They may be erect and flat or sometimes less open. The labellum is simple or very slightly lobed, usually very wide and showy without salient calli although normally showing more or less subtle keeled thickenings close to the base, usually of different colors; it is much larger and wider than the other segments, often flat but in M. candida embraces the column and in all species it is slightly fused to the column at their bases. The short column does not have a foot and presents two lateral auricles sometimes merged to each other through a fringe that surrounds the superior edge of the clinandrium. The anther is apical and bears two yellow hard pollinia. They possibly are pollinated by bees.[6]

Taxonomic notes

Miltonia flavescens, illustration.
This was the first Miltonia species to be described, originally classified under the genus Cyrtochilum, in 1834.

The first species to be described, among the ones today classified under the genus Miltonia, was originally published by John Lindley, in 1834, as Cyrtochilum flavescens. In this description Lindley notices that the flowers of this species turn orange color when drying and, for some confusion regarding the origin of the species, attributes it to Mexico instead of Brazil.[7] Two years later Lindley described another Miltonia species but, then, under the genus Oncidium, as O. russellianum in homage to Duke of Bedford. When describing this plant, Lindley considered it as a transition species pointing out that it was very different from the average Oncidium because of its purple colors and undivided lip.[8]

In 1837, Lindley received from Mr. Loddiges and from George Baker two other specimens of a very distinctive new species. Recognizing then this should in fact be a new genus, he proposed the name Miltonia to it as a homage to Lord Fitzwilliam Milton, an English orchid enthusiast. Lindley states then that the limits between a number of Oncidiinae genera, Cyrtochilum, Oncidium, Odondoglossum, Brassia and Miltonia, at that time classified as Vandaea, were yet to be perfectly established; although closely related, the differences should possibly be: Oncidium has a column with two ears and labellum distinctively lobed; Miltonia has a column with two ears and an entire labellum partially united to the column base; Odontoglossum and Cyrtochilum have winged columns and entire labelli but the former has it partially united to the column; and Brassia does not have any appendages on the column. It is interesting to notice that despite Lindley described the genus Aspasia in 1833, which is the most closely related to Miltonia, both by flower and vegetative morphologies, he did not mention it on Miltonia description.[9]

Three other Botanists were working with Miltonia species around the time Lindley described this genus. All recognized these plants should be classified under a new genus and, as communications were slower then, all proposed new genera: Knowles and Westcott also received also a plant of M. spectabilis and, just one month after Lindley, proposed for it the genus Macrochilus, calling the species Macrochilus fryanus;[10] the other one was Rafinescque who, in 1838, decided that the Oncidium russellianum already described by Lindley in 1836 should be under another genus and created for it the genus Gynizodon.[11] Both Macrochilus and Gynizodon are synonyms of Miltonia and no other species has ever been submitted to them.[2]

(CC) Photo: Dalton Holland Baptista
Miltonia russelliana
This is the less showy of Miltonia species because its sepals and petals do not really open, being always bent over the column, revealing only the lighter tip of its purple labellum.
(CC) Photo: Dalton Holland Baptista
Miltonia clowesii
This species has the same color pattern of M. russelliana, however, with a whiter labellum. On the other hand, M. clowesii flowers pointed segments are larger and wide opened making it look like a spider.

As Miltonia species are common plants, comparatively large, with also large flowers of bright colors, that, moreover, are spread mostly over an area of early settlements in Brazil all species but one were already described in 1850; six of them by Lindley, M. regnellii by Reichenbach[12] and M. moreliana by Achille Richard.[13] Despite the early description of M. moreliana in 1848, and two other as M. rosea by Lemaire in 1867,[14] and as M. warneri by George Nicholson in 1886,[15] Arthur Henfrey reduced it to a variety of Miltonia spectabilis in 1851,[16] and as such it was considered until 2002, when Cássio van den Berg reestablished it as a distinct species.[17] The last Miltonia species to be discovered was M. kayasimae, found by an orchid collector not far from the city of São Paulo, in an area around nine hundred meters of altitude nearby the top of Serra do Mar mountains. It was named after their collector by Guido Pabst in 1976. So far very few plants were found, all living at the same area.[18]

Since the genus Miltonia was established, many species, now classified under a number other genera, were submitted to it. The most noticeable cases were four of the five species of Miltoniopsis,[19] a genus proposed in 1889 but only really accepted in 1976.[20] Despite its somewhat similar flowers, Miltoniopsis are from cooler forests on the Andean slopes closely related to Cyrtochilum and only remotely related to Miltonia. Also five of the six Miltonioides species were occasionally considered as Miltonia until 1983 when Brieger and Lückel proposed this genus for them.[21] These are species of more delicate and narrower flowers, from Mexico and Central America, which some taxonomists claim might be better classified under the genus Oncidium to whom they are closely related.[22] The last common species which was occasionally classified under Miltonia is Chamaeleorchis warszewiczii,[23] which is related to Oncidium and some taxonomists identify as Oncidium fuscatum.

In 1983, Brieger and Lueckel, considering that four species of Miltonia, M. candida, M. cuneata, M. kayasimae and M. russelliana, show the junction of the labellum with the column in a different angle than the other species, proposed the genus Anneliesia for them.[24] Although this four species form a small sister clade to the rest of Miltonia species, the difference did not seem important enough to justify the acceptance of this new genus, therefore this proposal has not been generally accepted by the scientific community.[2]

In 2001, based on molecular analysis, Norris Williams and Mark Chase, transfered a species previously classified under the genus Oncidium, as O. phymatochilum, to Miltonia.[25] As this species shows a morphology that closer to Oncidium species than to Miltonia, because of its small yellowish flowers and highly branched inflorescence, this result and following transfer was a great surprise to most taxonomists. In 2005, Eric Christenson suggested a new genus and the name Phymatochilum brasiliense for it.[26] There is no consensus about the name to be generally accepted as yet.

Molecular analysis show that Miltonia most closely related genus is Phymatochilum and then Aspasia, Brassia and Ada, which are the most important genera included in this that is one of the eight clades that form the subtribus Oncidiinae of tribus Cymbidieae.[22]

Species

(CC) Photo: Dalton Holland Baptista
Miltonia cuneata
This is the hardest to grow among all Miltonia species, however, being all other very easy, this is not really a hard one. This species is the one that needs cooler conditions.
(CC) Photo: Dalton Holland Baptista
Miltonia kayasimae
This was the last species to be described and is the rarest one having being found just a couple of times in São Paulo State, in Brazil.

The species of Miltonia show many differences to each other and are very easy to identify, therefore, just the most evident differences are mentioned here; more details are given on individual species articles. The species are presented here according to their morphology and this order keeps no correspondence with phylogenetic relationships.

Regarding vegetative morphology Miltonia moreliana and Miltonia spectabilis can be immediately separated from the rest because their much flatter pseudobulbs, longer rhyzome and inflorescences completely covered by flattened bracts that bear only one highly flat flower. These are the species with largest flowers in the genus. They are closely related and usually are recognized because the flowers of M. moreliana usually have dark purple petals and sepals and the lip of a lighter bright purple while M. spectabilis has very light purple or white petals and sepals and a purple veined labellum, however, the real technical difference among the species is on the proportions of their segments which are much wider. Despite colors are often mentioned to identify species they are not accepted by taxonomy as enough to establish distinct species by themselves.[17] All other Miltonia species have similar vegetative appearance and only can be positively identified by their flowers.

Three species are very unique: Miltonia flavescens has the most narrow flowers, almost star shaped, with all segments of straw color with some purple bots on the base of petals and sepals which are more intense on the labellum almost forming stripes;[1] Miltonia candida is the only species with a labellum that embraces the column in a way that reminds the Cattleya species;[27] Miltonia russelliana is the less showy of Miltonia species because its sepals and petals do no really open, being always bent over the column, revealing only the lighter tip of its purple labellum.[8]

Miltonia regnellii shows the widest flower color variation among all Miltonia species; they can vary from white to yellow, pink and lilac with labelli also varying from white to dark purple. The flowers open in succession and slightly resemble the ones of M. spectabilis although much smaller. They actually are the Miltonia species with the smallest flowers.[12]

Miltonia kayasimae and Miltonia cuneata are somewhat similar and possibly are closely related, both have straw color petals and sepals almost entirely covered by large brown stains and white labelli, however, they show different proportions on the flowers segments. M. kayasimae has much wider petals and sepals and smaller labellum which, moreover, has a larger and more salient and complex entirely purple callus on its base which is delicate, more straight and simple, and just occasionally purple dotted on its apex on M. cuneata.[18]

Miltonia clowesii has the same color pattern of M. russelliana with light yellow greenish brown sepals and petals completely covered with large darker dots or stains and labellum of bright purple at the base and lighter apex, however here they are whiter. On the other hand, M. clowesii flowers pointed segments are larger and wide opened making it to remind a spider.[28]

Natural hybrids

(CC) Photo: Dalton Holland Baptista
Miltonia × bluntii
This is the natural hybrid of M. spectabilis and M. clowesii.

Considering its limited number of species, it is surprising that eight natural hybrids of Miltonia are currently known, a number that almost equals the number of species and also implies that the most important pollinator of the majority of the species possibly is the same. As the crossing of two species uses to produce variable plants most of these hybrids have been described more than once and some have three or four synonyms. M. spectabilis is the species which has produced the largest number of hybrids, five: Miltonia × bluntii when crossed with M. clowesii, Miltonia × cogniauxiae with M. regnellii, Miltonia × flava with M. flavescens, Miltonia × leucoglossa with M. candida and Miltonia × rosina with M. cuneata,[2] furthermore it is possible there is also one with M. moreliana, which has not yet been described because M. moreliana itself was earlier considered a variety of M. spectabilis.

M. candida, besides the hybrid already mentioned with M. spectabilis, also produced two others: Miltonia × binotii with M. regnellii and Miltonia × lamarckeana with M. clowesii.[2] The remaining hybrid, Miltonia × peetersiana used to be considered a synonym of M. × bluntii but because M. moreliana is now a species distinct from M. spectabilis it is its hybrid with M. clowesii, which has entirely purple flowers instead the one with light brown petals and sepals.[29]

Culture

Despite being easy to grow Miltonia species tent to be subject to spots on their thin leaves generally caused by fungi proliferation and normally, when exposed to the amount of light they need to a full bloom, their foliage gets a little bit too yellow colored, although they should never be exposed to full sunlight. Finding the right balance of light exposure to avoid the yellow leaves but still produce nice blooming is important and with some tentatives the grower will succeed. They are not much sensitive to temperature but it varies according to their origin M. cuneata being the one that grows cooler and M. moreliana the warmer grower all under intermediate temperature with at least 10ºC of variation between day and night. Despite they show a rest period after blooming, Miltonia always need to be watered, more abundantly during active growth. they need at least 65% of humidity and good ventilation all the time. Moderate weekly fertilizing with a balanced formula is beneficial during active growth. They may be potted in a compost of half-chopped Sphagnum, peat, and some medium sized lumps of charcoal, or mounted on plaques of vegetable fiber, however if mounted they will need more frequent waterings.[30]

References

  1. 1.0 1.1 Johnson, Andrés E. (2001). Miltonia flavescens in Las orquídeas del Parque Nacional Iguazú, 130. Fundación Vida Silvestre Argentina, Buenos Aires. ISBN 9509725412
  2. 2.0 2.1 2.2 2.3 2.4 Govaerts, Rafaël et al: World Checklist of Orchidaceae. The Board of Trustees of the Royal Botanic Gardens, Kew. Published on Internet (Access in March 2009).
  3. 3.0 3.1 Miller, David; Richard Warren; Izabel Moura Miller & Helmut Seehawer (2006). Miltonia in Serra dos Órgãos sua história e suas orquídeas, 323-6. Rio de Janeiro.
  4. Pabst, Guido & Dungs, Fritz (1978). Orchidaceae Brasilienses 2: 196, Brucke-Verlag Kurt Schmersow, Hildesheim. ISBN 3871050106
  5. Cogniaux, Célestin Alfred (1903). Miltonia in Flora Brasiliensis 3-6: 267-286. K.F.P. von Martius Ed.. Published on Internet, in Latin.
  6. Hoehne, Frederico C. (1940). Introduction in Flora Brasílica, Vol 12-1: 37-8. Secretaria de Agricultura de São Paulo.
  7. Lindley, John (1834). Cyrtochilum flavescens in Edwards's Botanical Register 19: t. 1627. England.
  8. 8.0 8.1 Lindley, John (1836). Oncidium russellianum in Edwards's Botanical Register 22: t. 1830. England.
  9. Lindley, John (1837). Miltonia spectabilis in Edwards's Botanical Register 23: t. 1992. England.
  10. Knowles, George B. & Westcott, Frederic (1837). Macrochilus fryanus in Floral Cabinet 1: 93. England.
  11. Rafinescque, Constantine S. (1838). Gynizodon russelliana in Flora Telluriana 4: 40..
  12. 12.0 12.1 Reichenbach, Heinrich G. (1850). Miltonia regnellii in Linnaea 22:: 851.
  13. Richard, Achille (1848). Miltonia moreliana in A.Rich., Portef. Hortic. 2: 38.
  14. Lemaire, Charles (1867). Miltonia rosea in Ill. Hort. 14.: t. 524.
  15. Nicholson, George (1886). Miltonia warneri in Ill. Dict. Gard. 2: 369.
  16. Henfrey, Arthur (1851). Miltonia spectabilis var. moreliana in Gard. Mag. 3:: 41. London.
  17. 17.0 17.1 Carlini-Garcia, L. A..; C. van den Berg; P. S. Martins (2002). A morphometric analysis of floral characters in Miltonia spectabilis and M. spectabilis var. moreliana. Lindleyana 17(3): 122-129. Published on Internet.
  18. 18.0 18.1 Pabst, Guido F.J. (1976). Miltonia kayasimae in Bradea 2: 88.
  19. Godefroy-Lebeuf, Alexandre. (1889). Miltoniopsis in Orchidophile 9: 63. Argenteuil.
  20. Garay, Leslie A. & Dunsterville, Galfrid C.K. (1976). Miltoniopsis in Venezuelan Orchids Illustrated. 6: 278.
  21. Brieger, Friedrich G. & Lückel, Emil (1983). Miltonioides in Orchidee 34: 130. Hamburg.
  22. 22.0 22.1 Williams NH, Chase MW, Fulcher T, Whitten WM (2001). Molecular systematics of the Oncidiinae based on evidence from four DNA sequence regions: expanded circumscriptions of Cyrtochilum, Erycina, Otoglossum, and Trichocentrum and a new genus (Orchidaceae) in Lindleyana 16(2): 113-139.
  23. Senghas, Karlheinz & Lückel, Emil (1997). Chamaeleorchis in Schlechter Orchideen I/C(33-36): 2305.
  24. Brieger, Friedrich G. & Lückel, Emil (1983). Anneliesia in Orchidee 34: 129. Hamburg.
  25. Williams, Norris H. & Chase Mark W. (2001). Miltonia phymatochila in Lindleyana 16: 284.
  26. Christenson, Eric A. (2005). Phymatochilum brasiliense in Richardiana 5: 195.
  27. Lindley, John (1838). Miltonia candida in Edwards's Botanical Register 24 (Misc.): 25. England.
  28. Lindley, John (1840). Miltonia clowesii in Sertum Orchididacearum: t. 34. England.
  29. Reichenbach, Heinrich G. (1886). Miltonia peetersiana in Gardener Chronicles, n.s., 26: 326.
  30. Baker, Charles O & Baker, Margaret L. (2006), Miltonia in Orchid Species Culture Oncidium/Odontoglossum Alliance: 340-7, Timber Press. ISBN 9780881927757
 (Read more...)
Caesar Schinas 09:25, 8 June 2009 (UTC); Meg Ireland 08:00, 18 June 2009 (UTC); Howard C. Berkowitz 14:32, 5 June 2009 (UTC) Daniel Mietchen 17:24, 4 June 2009 (UTC) 5


Developed Article Scuticaria: A genus of orchids, closely related to Bifrenaria, formed by nine showy species of cylindrical leaves, which exist in three isolated areas of South America. [e]

Scuticaria
Scuticaria steelei
Scuticaria steelei
Scientific classification
Domain: Eukaryota
Kingdom: Plantae
Division: Magnoliophyta
Class: Liliopsida
Order: Asparagales
Family: Orchidaceae
Subfamily: Epidendroideae
Tribe: Cymbidieae
Subtribe: Maxillariinae
Genus: Scuticaria
Lindl. 1843
Type species
Maxillaria steelei
Hook. 1837
Species
Synonyms
  • None

Scuticaria is a genus in the orchid family formed by nine species of showy flowers and long cylindrical leaves. They are epiphytic, occasionally lithophytic or terrestrial, that grow pending and are cespitosus, or reptant and ascending, which exist in three isolated areas of South America, the Amazon Forest in Ecuador and the Serra do Mar and Serra da Mantiqueira mountains in Brazil, both in shady and sunny places.

The genus Scuticaria has been traditionally placed close to Maxillaria but recent research shows they are more closely related to the genus Bifrenaria. Despite their interesting appearance, the are hardly seen in nature and, because they culture is complicated, they are not common in private collections and orchid shows either. No other use for these species is reported besides ornamentation. Because it is a well established genus, formed by few species that are reasonably easy to separate, there were few publications about them during the last decades.

Distribution and habit

Despite there are few species, Scuticaria inhabit varied climates, disperse in a very uneven way though all countries of South America northern to Bolivia, this excluded, and also in areas of Mata Atlântica in Brazilian Southeast. No species is common in nature, being just occasionally or even rarely found.

The species with wider range is Scuticaria steelei which inhabits open clearings at higher elevations of central Amazon, jungles known as matas de terra firme, up to eight hundred meters of altitude.[1] Although this species occupies wide area, it is not found very often.[2] Another species from Amazon, however, in a much more restricted area, just in Guyana, in places where the altitude is lower and the humidity is higher, is Scuticaria hadwenii var. dogsonii.[3]

Endemic in another area of Amazon, separated but not that far from the habitat of Scuticaria steelei, on southeastern Ecuador, close to the place where the Andes starts, in humid and slightly colder forests, on the mountains up to 1,300 meters of altitude, it is found Scuticaria salesiana.[4] Under the same conditions but in wider areas, that encompass the southeast of Ecuador and northeast of Peru, lives S. peruviana.[5] All species from Amazon are always epiphytic.

The remaining species inhabit the area occupied by Brazilian Atlantic Forest. The only species that can be found widespread through several states is Scuticaria hadwenii, in the humid jungles of Serra do Mar from Santa Catarina to Bahia States,[6] generally found living epiphytic at middle height over thick tree stems.[7] Other species occasionally found, although often under living litophytic over rocks and gatherings of fallen leaves in sunny areas of the mountains of São Paulo and Rio de Janeiro, is S. strictifolia.[8]

Scuticaria irwiniana, second and last rupicolous species, exists only on the mountains of Minas Gerais State, found in sunny or shadier places up to two thousand meters of altitude.[9] Two are the species from Espírito Santo State, S. novaesii and S. kautskyi, both endemic of restricted areas in the dry jungles of the countryside.[10] The last Scuticaria species is S. itirapinensis, which has been found only a couple of times in the west-central dry woods of São Paulo State, in an area which has been highly deforested, close to Itirapina. There are no records or reports on this species, both in nature and under culture, during the last twenty five years. It is speculated about the possibility of its extinction.[11]

Description

(CC) Photo: Dalton Holland Baptista
Scuticaria novaesii.
This is an endemic species from Espirito Santo State in Brazil discovered only in 1981.

The species subordinated to genus Scuticaria are characterized by being plants of thick cylindrical roots covered by thick vellamen. Their stem is formed by a ordinarily short rhizome, slightly elongated in some species; and by cylindrical almost inconspicuous pseudobulbs of the same diameter or slightly thicker than the unique leaf born on their apexes, because they generally are covered by small dried scaling steaths. The leaves may be erect or pending up to one meter long. The inflorescences grow from the said steaths and almost always bear just one flower, exceptionally two in one species, and always is much longer than the pseudobulbs, bearing showy yellow, orange, purple or greenish flowers, with petals and sepals plain, stained or striped, usually by light brown but also by diverse combinations and shades of the other mentioned colors. Ordinarily the labellum presents contrasting colors, frequently with white areas.[8]

The flowers are large, wide open, and last during about two weeks.[2] They have sepals of similar sizes and form an almost invisible chin with the column foot. The petals may be similar to the sepals but smaller, or much smaller and with a much narrower base, occasionally showing different patterns or colors. The labellum articulates with the column, is trilobed, with comparatively small lateral lobes and larger terminal, which has variable shapes with diverse patterns and a callus under to column. The later is é semi-cylindrical, slightly arching, erect and thick, without any kind of appendix, ending in an apical anther and elongated in a small foot at the base. The flowers bear to pairs of pollinia of different sizes. The caudicle is narrow and the retinacle is small. The fruits resemble the ones of Maxillaria.[8] There are no observation records of pollinators activities but Scuticaria are supposedly pollinated by Euglossini bees.[7]

Taxonomic notes

Scuticaria steelei
Illustration of original description published by William Jackson Hooker in 1837.

In May of 1837, the English Botanist William Jackson Hooker received a drawing and a dried sample of a plant, sent by an orchid grower from Liverpool, together with a note explaining that the plant arrived from Demerara, in Guyana, in July of the preceding year. Hooker described this species, classifying it under the genus Maxillaria, calling it M. steelei, in homage to its discover. In his description, Hooker affirms that the plant is highly interesting and an excellent addition to the known epiphytic species because it shows cylindrical leaves almost one meter long, different from anything ever found.[12] few months later, John Lindley published again the species Hooker described, however, adding more information. Two years earlier, several plants had been sent from Demerara and Lindey reports that he had previously informally classified this species as Maxillaria flabellifera which, under this name, could be found in several orchid collection in England. Because he had not yet described this species, he accepts the priority of the name chosen by Hooker. Lindley adds, however, that he had some doubts about the classification, of a species so different from any other known so far, under the genus Maxillaria.[13]

In 1843, Lindley published a revision of a group of orchids classified as tribus Maxillaridae, then subordinated to Vandeae, a subfamily of Orchidaceae at the time. In this revision, he indicates that much work is needed till the limits between each genus within this tribus can be established and states his doubts regarding some of the new genera he was proposing, despite being very sure of other ones. Lindley suggested the division of Maxillaridae in twenty five genera, being Scuticaria one of the genera he considered well established. When describing this new genus, Lindley based on morphologic characteristics of Maxillaria steelei Hook., selected as the Type species of Scuticaria with the name Scuticaria steelei.[14] This name comes from Latin scutica, flagellum, in reference to the long cylindrical leaves that the species of this genus show, similar to the leather whips used to punish.[8]

Strangely, because he published the genus Scuticaria many years earlier, in 1851, Lindley described another species now considered part of this genus, classifying it under Bifrenaria. It is speculated that possibly because it was found in Brazil on the same area in the southeast where most of Bifrenaria were common, or because he believed that two species separated by so long distance belonged to the same genus. It was Scuticaria hadwenii.[15] Few months later, Jules Émile Planchon corrected Lindley moving it to the genus where it is subordinated today.[16] In 1851, the only two common Scuticaria species were described and the genus well established, therefore no later confusion about the classification of any species subordinated to this genus ever happened.[17]

Almost one century passed before any important new information were published. In 1881 Heinrich Gustav Reichenbach described Scuticaria dogsonii, originated from Guyana,[3] but in 1892, Berthold Stein, considering that the only difference it shows from Scuticaria hadwenii is the fact it bears two flowers each inflorescence, reduced it to a variety of the later.[18] In 1903, Célestin Alfred Cogniaux, when revising all orchids species from Brazil, cites two other varieties of Scuticaria hadwenii which, because just show color differences, can not be accepted as such today.[19] Finally, in 1947, Frederico Carlos Hoehne described a new species, Scuticaria strictifolia, yet similar to Scuticaria hadwenii, although showing some slight differences on the labellum structure, besides their normally lithophytic habit and erect leaves.[20]

(CC) Photo: Dalton Holland Baptista
Scuticaria irwiniana.
This is one of the two Scuticaria species with erect leaves and the one with the shortest leaves among all species.

If few species were known so far, after 1968 the number of described species triplicated. All species described during the later years are uncommon and inhabit restricted areas, some are very rare or even supposedly exctinct. In 1968 Robert Louis Dressler described Scuticaria salesiana, discovered in Ecuador in an area far apart from the other Scuticaria range.[4] In 1972, Guido Pabst described Scuticaria kautskyi, found in Espírito Santo State, [21] in southeast Brazil and, during the following year, published two species at once, S. itirapinensis and S. irwiniana.[22] In 1982, other species was discovered in Espírito Santo, Scuticaria novaesii.[10] The last described species was S. peruviana, found in Peru in 2002, in the same region of S. salesiana, to which it is related.[5]

Despite Lindley indicated the possibility of Scuticaria being closely related to Bifrenaria when he initially described S. hadwenii under this genus, all later taxonomists always included Scuticaria on the same group Maxillaria were.[23] It was just in 2000 that the first proofs of Scuticaria closer proximity to Bifrenaria started being published.[24] In 2002, a detailed research about the phylogeny of Bifrenaria performed molecular analyses on two Scuticaria species while chosing them as out groups. This study claims that the phylogenethic internal relationships among Scuticaria species so far remain unknown.[25]

It is known that other orchid genera bearing cylindrical leaves devolved this sort of leaves as a defense to climate changes their habitats were going through along the eras. Terete leaves are capable of much more water and nutrients and to face longer drought periods than species bearing thin leaves, on the other hand, almost all epiphytic species presenting the former type of leaves show more or less atrophied pseudobulbs since the leaves carry on its accumulating role. It is a supposition that Scuticaria species should have once inhabited much drier through their evolution. Because most of the species are found in shadier and more humid species now, this may one of the reasons why their culture uses to be complicated, possibly because the delicate balance they reached in nature is broken. For the same reason it is supposed their frequency in nature is only occasion or rare.[7]

Species

(CC) Photo: Dalton Holland Baptista
Scuticaria hadwenii
This is the most variable among "Scuticaria species and widespead through a large area in Southeast of Brazil.

Because of its highly particular morphologic characteristics which allow immediate identification, their restricted species distribution, and their comparatively low variability, since the genus Scuticaria was established by Lindley, only then species were formally described and it has never been great confusion separating each species. Form these ten, nine are generally accepted, the tenth being ordinarily considered a variety, and, under this condition, also accepted.[17] For identification purposes, the species can be split as follow:

Only two species present erect leaves and are the only ones frequently found as lithophytes, Scuticaria irwiniana, easily known recognized because of its flowers without any stains on the internally entirely purple and externally whitish sepals and petals, with white labellum, striped of purple. This species generally can be identified even without flowers because of its reptant, slightly ascendant growth, and longer rhyzome than any other species.[22] Scuticaria strictifolia also has erect leaves but occasionally, when cultivated under insufficient light, their leaves can be narrower and slightly bent making the distracted observer find hard to differentiate it from S. hadwenii.[8] The Brazilian taxonomist Guido Pabst considered this species a variety of the later.[9]

(CC) Photo: Dalton Holland Baptista
Scuticaria strictifolia
It is very close to Scuticaria hadwenii but has erect leaves; different colors and diverse callus structure on the labellum.

All species remaining are ephiphytic with pendent habit. Scuticaria hadwenii, due to its several more or less isolated groups of populations along Serra do Mar, mostly on the west side of this chain of mountains, spreading throughout the interior highland in some states of Brazil, is the Scuticaria species that presents most variable colors.[8] It can be separated from S. strictifolia because shows leaves always pending, flowers of more vivid colors and by the interior of the labellum, which ordinarily is more pubescent. There is a variety denominated dogsonii, native from Guyana, which is more floriferous.[3]

The two other species from Espírito Santo State are highly different to each other. Scuticaria kautskyi usually has more or less uniform orange color on its sepals and petals, with their bases slightly lighter and dotted of greenish yellow. Their labellum is white showing few colored drawings and narrow terminal lobe, slightly deflected.[21] The other species from this state, Scuticaria novaesii presents flowers with green-yellow segments, intensely spotted with dark brown and wide and flat labellum terminal lobe, with clearly marked by radial multicolored lines.[10]

Scuticaria itirapinensis, the last species of brazilian southeast, is the one that closely resembles Amazonian Scuticaria steelei, although it can be easily separeted because of its strong yellow flowers and much shorter leaves, besides slight differences on the proportions of floral structures.[22] A Scuticaria steelei presents entirely pale yellow flowers, completely covered by spaced small darker stains, However in is not even necessary to observe the flowers to identify it as their leaves are about one meter long, and there are references of plants measuring almost one and a half meter.[2]

The last two Scuticariaare isolated in forests of Peru and Ecuador and are similar to each other. They are different from all other because of the proportions of floral segments. The labellum is much larger when compared to their sepals and petals than it is on other species. Moreover, their petals are striped of brown and much smaller than the sepals, showing a greater difference than it is found on the other species. From each other, they can be separated mostly by the shape of the labellum. Scuticaria salesiana presents more rounded intermediate lobe, and Scuticaria peruviana has it more rectangular, with the apex truncated, almost in a straight line.[5]

In 2008, a new species of Scuticaria, S. bahiensis has been described from Bahia state in Brazil but so far it remains mostly unknown.[26]

Culture

Scuticaria itirapinensis.
This showy species has been found just a couple ot times in nature and has not been found or seen anywhere during the last decades.

In his book Flora Brasilica, the Brazilian Botanist Frederico Hoehne strongly recommended the culture of Scuticaria species because of their beautiful flowers and interesting vegetation, however, soon later he admits that all species then cultivated by São Paulo Botanic Garden had died after two or three years. Indeed, he claims that to successfully grow them a special environment needs to be created.[8] These plants are not easy to maintain under culture. Only recently, with the help of modern technology, timers and foggers that keep the humidity constant, the growers have been finally capable of keeping them out of their natural environment for several years.

There are four different sorts of culture according to the origin of each species. S. steelei and S. hadwenii var. dogsonii are the species that need higher temperature and humidity. The two rupicolous species, S. irwiniana and S. strictifolia ate the ones which need more light and constant ventilation besides drier culture conditions.[22] S. peruviana and S. salesiana take slightly cooler temperatures than the other species although still need humidity mostly during the early morning hours.[5] The other species need less light than the mentioned ones. All species should be preferably mounted on plaques of vegetable fibers because of their pending habit, the rupicolous species may alternatively potted in well drained pots. Scuticaria are delicate plants that like to remain untouched during several years because their roots easily resent on replants.[7]

References

  1. Miranda, Francisco: Orquídeas da Amazônia Brasileira, pp. 43. Ed. Expressão e Cultura, 1996. ISBN 8520802087
  2. 2.0 2.1 2.2 Freitas Luz, Francisco J.: Orquídeas na Amazônia, pp. 59. Instituto Brasileiro de Cultura, Ed. On Line, 2001. ISBN 8520802087
  3. 3.0 3.1 3.2 Reichenbach, Heinrich Gustav: Scuticaria dogsonii in Gardeners' Chronicle vol.15: pp.9. London, 1881.
  4. 4.0 4.1 Dressler, Robert Louis: Scuticaria salesiana in Orquideologia vol.3(2): pp.3. Revista de la Sociedad Colombiana de Orquideologia. Medellin, 1968.
  5. 5.0 5.1 5.2 5.3 Bennett, David E. & Christenson, Eric: Scuticaria peruviana in Orchid Digest 66: pp.64. Berkeley, California, 2002.
  6. Toscano de Brito, Antônio & Cribb, Phillip: Orquídeas da Chapada Diamantina, pp. 284. Ed. Nova Fronteira, 2005. ISBN 8520917828
  7. 7.0 7.1 7.2 7.3 Miller, David; Richard Warren; Izabel Moura Miller & Helmut Seehawer: Serra dos Órgãos sua história e suas orquídeas, pp. 294. Rio de Janeiro, 2006.
  8. 8.0 8.1 8.2 8.3 8.4 8.5 8.6 Hoehne, Frederico Carlos: Scuticaria in Flora Brasilica, vol. 12-7 pp.342. Instituto de Botânica de São Paulo, 1953.
  9. 9.0 9.1 Guido Pabst & Fritz Dungs: Orchidaceae Brasilienses vol. 2 pp. 187, Brucke-Verlag Kurt Schmersow, Hildesheim, 1977. ISBN 3871050107
  10. 10.0 10.1 10.2 Barros, Fábio & Catharino, Eduardo L.M.: Scuticaria novaesii, nova espécie de Orchidaceae do Brasil. Hoehnea vol. 9: pp. 52-62, São Paulo, 1982.
  11. Records of species displayed during orchid shows. Archives of Coordenadoria das Associações Orquidófilas do Brasil - CAOB. Accessed October 2008.
  12. Hooker, William Jackson: Scuticaria in The Botanical magazine 64: t. 3573. Ed. William Curtis, London, 1837. Published on Internet.
  13. Lindley, John: Scuticaria steelei in Edward's Botanical Register Vol.2, t.1986. James Ridgway & Sons Ed. London, 1837. Published on Internet.
  14. Lindley, John: Scuticaria in Edward's Botanical Register Vol.29 (miscelanea), pp.14. James Ridgway & Sons Ed. London, 1843. Published on Internet.
  15. Lindley, John: Bifrenaria hadwenii in Paxton's Flower Garden. John Paxton Ed., London ,1851.
  16. Planchon, Jules Émile: Scuticaria hadwenii in Flore des Serres et des Jardins de l'Europe vol.7: pp.239. Ghent, 1852.
  17. 17.0 17.1 R. Govaerts, M.A. Campacci (Brazil, 2005), D. Holland Baptista (Brazil, 2005), P.Cribb (K, 2003), Alex George (K, 2003), K.Kreuz (2004, Europe), J.Wood (K, 2003, Europe) (Novembro 2008). World Checklist of Orchidaceae. The Board of Trustees of the Royal Botanic Gardens, Kew. Published on Internet. (Accessed March 2009).
  18. Stein, Berthold: Scuticaria hadwenii var. dogsonii in Orchideenbuch, 1892.
  19. Cogniaux, Célestin Alfred: Scuticaria in Flora Brasiliensis Vol.3 Part.6: pag. 78-81. K.F.P. von Martius Ed., 1903. Publishe on Internet.
  20. Hoehne, Frederico Carlos: Scuticaria strictifolia in Arquivos de Botânica do Estado de Sao Paulo, n.s, f.n.2, pp.88. Instituto de Botânica de São Paulo, 1947.
  21. 21.0 21.1 Pabst, Guido F.J.: Scuticaria kautskyi in Bradea 1: pp.169 Boletim do Herbarium Bradeanum. Rio de Janeiro, 1972.
  22. 22.0 22.1 22.2 22.3 Pabst, Guido F.J.: Scuticaria irwiniana and Scuticaria itirapinensis in Bradea 1: pp.336-7 Boletim do Herbarium Bradeanum. Rio de Janeiro, 1973.
  23. Dressler, Robert Louis: Phylogeny and classification of the orchid family. Cambridge University Press, 1993.
  24. Whitten, W. Mark; Williams, Norris H. & Chase, Mark W.: Subtribal and generic relationships of Maxillarieae (Orchidaceae) with emphasis on Stanhopeinae: combined molecular evidence. American Journal of Botany. 2000;87: pp. 1842-1856, 2000. published on Internet.
  25. Koehler, Samantha: Estudo taxonômico e análise cladística do complexo Bifrenaria Lindl. (Maxillarieae, Orchidaceae). Universidade Estadual de Campinas. Instituto de Biologia, December 2001. published on Internet.
  26. K.L.Davies & Stpiczynska. (2008). Scuticaria bahiensis in Orchid Review 116: 344.
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Caesar Schinas 09:25, 8 June 2009 (UTC); Meg Ireland 08:00, 18 June 2009 (UTC) Daniel Mietchen 17:24, 4 June 2009 (UTC) 4


Torture: Add brief definition or description
Torture (Read more...)
Milton Beychok 01:33, 19 June 2009 (UTC); Drew R. Smith 03:58, 19 June 2009 (UTC); Peter Schmitt 23:23, 24 June 2009 (UTC) Howard C. Berkowitz 14:32, 5 June 2009 (UTC) 5


Developed Article Milpa agriculture: A form of swidden agriculture that is practiced in Mesoamerica. Traditionally, a "milpa" plot is planted with maize, beans, and squash. [e]

(CC) Photo: Joey Hipolito
Milpa plots are found from Chihuahua to Central America. This one is in Oaxaca, Mexico. Squashes are being grown between the rows of maize.

Milpa agriculture is a form of swidden agriculture that is practiced in Mesoamerica. Traditionally, a "milpa" plot (from the Nahuatl word for "corn field") is planted with maize, beans, and squash (known as the Three Sisters) and might include a variety of other plants. These plots are planted for two or three years and then allowed to lie fallow for some years in order to restore the fertility of the soil. Milpa agriculture varies somewhat by region and it has changed in a variety of ways in different areas but it remains an important part of life for millions of people throughout Mesoamerica.

Maize and beans

Maize and beans, which are the staples of the Mesoamerican diet, complement each other in terms of the health of the fields as well as the health of the people who eat them. Maize requires high levels of nitrogen in the soil to grow properly and quickly depletes the soil if planted alone. Bean plants (genus Phaseolus), on the other hand, are high in nitrogen and their presence extends the life of the maize plot significantly by helping to keep nitrogen levels healthy. One might say that the maize repays the debt it owes to the beans by providing stalks for the bean plants to cling to as they grow. Squashes, generally grown between the rows of maize stalks, also figure into this symbiotic relationship, as they cover the ground in between the rows of corn and help to keep weeds down.

Maize and beans also compliment each other nutritionally. If eaten alone, one would need to consume large amounts to fulfill human dietary requirements but when eaten together, they achieve "protein complimentarity."[1] That is to say, a person needs less total food if the two are eaten together. This diet is further supplemented by several varieties of squash and other foods that are planted along side the maize and beans. In many areas, the "Mesoamerican trio" (maize, beans and squash) is complemented by wild or semi-domesticated plants that grow in and around the milpa.

(CC) Photo: Lorena Pajares
Harvesting beans from a mountainside milpa in Chiapas, Mexico.

Planting and harvesting

Milpas are traditionally cultivated using the swidden, or "slash and burn" system. The forest is cut, allowed to dry and then burned. The left over ashes are then mixed into the soil as a fertilizer. Maize and beans are planted together in the same hole while squash is planted separately, in between the rows of maize. Other cultigens may be planted in a separate section of the field or scattered among the maize.

The fields must be weeded several times throughout out the season. This is backbreaking work, especially in areas where a milpa is likely to be planted on the slope of a mountain several hours walk from home and it is usually done mostly by hand with the help of machetes and hoes. Later in the season, weeding is less needed, as the broad leaves of the squash plants keep unwanted plants to a minimum and the maize grows out of reach of the weeds.

Beans and squash are harvested as they ripen. As the maize ripens, a few elotes (or fresh cobs) may be picked to be eaten right away but most of the plants are bent near the top and the cobs are allowed to dry in the field. Once they are dry, they are collected and stored for later use in traditional Mesoamerican foods like tortillas and tamales. The dry stalks are often collected and sold at market as fodder for animals and the husks are frequently saved for use in preparing tamales.

The weeds and left over corn stalks from the previous season will be burned to prepare the fields for planting for one or two more years but then they will be left to fallow for as much as ten or more years. This practice has been abandoned in many areas because of the shortage of land that is available to many families. Instead, animal dung or ashes from the hearth at home may be added to those mixed into the soil and many farmers have turned to industrial fertilizers that allow them to cultivate the same plot year after year.

The ritual life of the milpa

One of many ornate murals in the Zapatista community of Oventic in Chiapas, Mexico. One need not look far to discover the intimate connection between maize and people in Mesoamerican culture. Here, a human head wearing the characteristic Zapatista ski mask appears on every kernel.

Several important rituals are performed at key points during the growing season. The timing of these rituals varies by region and altitude due to variations in climate and the length of the growing season but they are invariably tied to significant events in the ritual calendar. Among the Maya, agricultural rituals are performed at the full moon[2] while in central Mexico, each ceremony is associated with one of the eighteen months in the solar calendar and occur at periods of twenty days.[3] Ceremonies are performed at each major stage in the development of the maize, beginning with the planting and concluding with the harvest.

Agricultural rituals also vary in form according to local traditions, but common themes are found throughout Mesoamerica. One such theme is the ritual reenactment of the creation of the world by marking the four cardinal directions. Among the K'iche' of highland Guatemala, offerings are made at the four corners of the milpa. The Kekchi plant maize in the center of the field first and then plant in each of the fours directions.[2] In Tepotzlán, the Nahuas place crosses made from pericón at the four corners of the field.[3] And in the Sierra Madre of northern Mexico, the Tarahumara open their work parties by scattering tesguino, or corn beer, to the four directions.[4][5] Other themes include sacrifices and offerings to indigenous deities or Catholic saints and prayers for rain.

Maize also figures prominently in ritual activities that are not directly connected to the cultivation of the milpa. In Amatlán, Nahua villagers say, "Corn is our blood,"[6] meaning, "corn defines our way of life" in everything from ethnic identity to every day activities. In fact, the Popul Vuh reveals that the first mothers and fathers of the Maya were actually formed from maize.[7] Maize, and especially the intimate connection between maize and humans, arises again and again in both ancient and modern myths.

Modern trends

One consequence of the changes in land ownership patterns introduced through Spanish colonial society and some of the agrarian reforms carried out by the modern states in the region is that many families do not own enough land to produce all of the food that they need in a year. Consequently, the practice of fallowing the fields has disappeared or nearly disappeared in many areas. Today, many farmers use chemical fertilizers to increase their fields’ production and engage in other economic activities to earn enough money to provide for their families.

Even when farmers have access to enough land to feed their own families, they are often unable to market their surplus. Trade liberalization and market deregulation policies adopted throughout the region in the 1980s and 1990s have meant that maize and other staple crops produced on a very large scale by U.S. corporations are frequently available at lower prices in local markets than the products of local farmers.[8] Thus, many farmers have also converted their fields to other crops. Today, cash crops destined for export to the U.S. include broccoli, snow peas, cucumbers, Brussels sprouts, and carnations.

Despite the market pressures felt by small scale farmers in Mesoamerica to turn away from traditional agriculture, milpas may hold information that can help improve modern agricultural methods. Monocrop agriculture creates artificial growing conditions that are significantly less biologically diverse than the natural ecosystems that they replace. This results in rapid depletion of the soil, which is generally counteracted by the application of chemical fertilizers that return nutrients to the soil but may have damaging effects in the long-term. Though it is unlikely that the balancing diversity of milpa agriculture can be reproduced on an industrial scale, the indigenous knowledge embedded in its design may provide some guidelines for improving industrial techniques. "By studying [the milpa's] essential features," writes Charles C. Mann, "researchers may be able to smooth the rough ecological edges of conventional agriculture."[9]

Guatemala landscape.JPG

Notes

  1. Edward F. Fischer and Carol Hendrickson. 2003. Tecpán Guatemala: A Modern Maya Town in Global and Local Context. ISBN 0-8133-3722-4
  2. 2.0 2.1 Karen Bassie-Sweet. 1999. "Corn Deities and the Complementary Male/Female Principle". Paper presented at La Tercera Mesa Redonda de Palenque.
  3. 3.0 3.1 Thomas L. Grigsby and Carmen Cook de Leonard. 1992. "Xilonen in Tepoztlan: A Comparison of Tepoztecan and Aztec Agrarian Ritual Schedules". Ethnohistory 39(2):108-147. DOI:10.2307/482390
  4. John G. Kennedy. 1963. "Tesguino Complex: The Role of Beer in Tarahumara Culture." American Anthropologist 65(3):620-640. Pp. 623.
  5. Jerome M. Levi. 1999. "Hidden Transcripts among the Rarámuri: Culture, Resistance, and Interethnic Relations in Northern Mexico." American Ethnologist 26(1):90-113. Pp. 101.
  6. Alan R. Sandstrom. 1991. Corn Is Our Blood: Culture and Ethnic Identity in a Contemporary Aztec Indian Village. ISBN 0806123990
  7. Dennis Tedlock. 1996. Popol Vuh: The Mayan Book of the Dawn of Life. ISBN 0684818450
  8. John Weeks. 1999. Trade liberalization, market deregulation and agricultural performance in Central America. Journal of Development Studies 35(5): 48-75. DOI: 10.1080/00220389908422591
  9. Charles C. Mann. 2005. 1491: New Revelations of the Americas Before Columbus. New York: Vintage Books. Pp. 221.
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Milton Beychok 00:15, 29 May 2009 (UTC);
Arne Eickenberg 14:15, 17 June 2009 (UTC); Peter Schmitt 23:23, 24 June 2009 (UTC)
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Miltonia
Miltonia spectabilis
Miltonia spectabilis
Scientific classification
Domain: Eukaryota
Kingdom: Plantae
Division: Magnoliophyta
Class: Liliopsida
Order: Asparagales
Family: Orchidaceae
Subfamily: Epidendroideae
Tribe: Cymbidieae
Subtribe: Oncidiinae
Genus: Miltonia
Lindl. 1837
Type species
Miltonia spectabilis
Lindl. 1837
Species
Synonyms

Miltonia is an orchid genus formed by nine epiphyte species and eight natural hybrids inhabitants of Brazilian Atlantic Forest, one species reaching the northeast of Argentina and east of Paraguay. This genus was established by John Lindley in 1837, when he described its type species, Miltonia spectabilis. Many species were attributed to Miltonia in the past, however, today, the species from Central America and from cooler areas on northwest of South America have been moved to other genera. Miltonia species have large and long lasting flowers, often in inflorescences with several of them. This fact, allied to being species that are easy to grow and to identify, make them a favorite of orchid collectors all over the world. Species of this genus are extensively used to produce artificial hybrids.

Despite the fact that Miltonia is now a well established genus, most of its species were originally classified under other genera as Cyrtochilum, Oncidium, Odondoglossum, and Brassia. All were discovered between 1834 and 1850 with the exception of M. kayasimae, discovered only in 1976.

Distribution

(CC) Photo: Dalton Holland Baptista
Miltonia candida
This is the only species with a labellum that embraces the column in a way that reminds Cattleya species.

Miltonia species range starts on the area of Missiones in the northeast of Argentina[1] and east of Paraguay[2] and spreads north along the Brazilian mountains of Serra do Mar and its branches up to the State of Pernambuco on Brazilian northeast. They occupy mostly areas between 200 and 1,500 meters of altitude meters, however the majority of the species are more often found about 600 to 900 meters. Miltonia species can be found from shady areas inside the forest to areas more exposed to the sun, however never are under full sunlight; usually in ventilated places where they receive plenty humidity during the night and early morning. They are always epiphyte and, because they grow very fast, each pseudobulb originating two new growths every year, they soon form large colonies.[3]

Miltonia russelliana and M. flavescens are the ones with the widest dispersion and found at lower altitudes. M. flavescens is the only species that exists in countries other than Brazil and is also the one that spreads farther north. M russelliana range starts on Rio Grande do Sul and ends at Bahia State. M. regnellii is also widespread although does not go northern than Rio de Janeiro. M. moreliana is a species more common at lower altitudes and warmer areas existing from Rio to Pernambuco. Miltonia candida, M. clowesii and M. spectabilis are restricted to the four states of Region Southeast of Brazil. Miltonia cuneata is just from São Paulo and Rio and the one that grow at highest altitudes. M. kayasimae is the only species really rare; it has been found just a couple of times in a very restricted area close to Salesópolis, in São Paulo State. The mountains area between São Paulo and Rio de Janeiro, where almost all species do exist may be considered the center of distribution of Miltonia.[4]

Description

(CC) Photo: Dalton Holland Baptista
Miltonia regnellii
This species shows the widest flower color variation among Miltonia species; they can vary from white to yellow, pink and lilac with labelli also from white to dark purple.

Miltonia are comparatively medium large orchid plants reaching about fifty centimeters height. They present subcaespitous growth, that means their pseudobulbs are not tightly packed but slightly spaced by a rhyzome, that is longer than on caespitous plants, with length between two and five centimeters. Their roots grow along the rhyzome in high numbers. They are white, comparatively thin, usually short and hardly branched. The rhyzome is covered by dried imbricating steaths which get increasingly larger at the base of pseudobulb becoming articulated foliar steaths that partially cover them. The pseudobulbs and leaves vary in color from yellowish bright light green to olive green depending on the species and to the amount of sunlight they are exposed to. They may be more oval and laterally highly flattened to slightly tetragonal and elongated and almost always bear two apical leaves. The leaves are narrow, flexible and hardly larger than three centimeters wide and forty long with the apexes rounded sometimes slightly pointed. Some species are about half of this size. The inflorescences are one or two per pseudobulb, shoot from their bases behind the protecting steaths. They are erect and never branched, often longer than the leaves, bearing from one to twelve moderately spaced flowers that open at the same time or in succession holding three or four opened all the time, when the older fades a new one opens. The older flowers of species with white lips that open in succession usually get yellower about the time the next flower opens although they still last one more week before fading. The first to bloom is M. cuneata, during late winter, but the majority of species bloom from late spring to late summer.[5]

The flowers of Miltonia vary from four to fifteen centimeters across; the larger are the ones with fewer flowers. Their colors vary from entirely white and pink to dark purple, pale yellow or lilac when plain, or they may highly spotted but then usually they are greenish or brownish with a contrasting labellum often white with purple dots, stains or veins close to the base.[3] The petals and sepals shapes are highly variable from species to species but always somewhat similar to each other within a species. They may be erect and flat or sometimes less open. The labellum is simple or very slightly lobed, usually very wide and showy without salient calli although normally showing more or less subtle keeled thickenings close to the base, usually of different colors; it is much larger and wider than the other segments, often flat but in M. candida embraces the column and in all species it is slightly fused to the column at their bases. The short column does not have a foot and presents two lateral auricles sometimes merged to each other through a fringe that surrounds the superior edge of the clinandrium. The anther is apical and bears two yellow hard pollinia. They possibly are pollinated by bees.[6]

Taxonomic notes

Miltonia flavescens, illustration.
This was the first Miltonia species to be described, originally classified under the genus Cyrtochilum, in 1834.

The first species to be described, among the ones today classified under the genus Miltonia, was originally published by John Lindley, in 1834, as Cyrtochilum flavescens. In this description Lindley notices that the flowers of this species turn orange color when drying and, for some confusion regarding the origin of the species, attributes it to Mexico instead of Brazil.[7] Two years later Lindley described another Miltonia species but, then, under the genus Oncidium, as O. russellianum in homage to Duke of Bedford. When describing this plant, Lindley considered it as a transition species pointing out that it was very different from the average Oncidium because of its purple colors and undivided lip.[8]

In 1837, Lindley received from Mr. Loddiges and from George Baker two other specimens of a very distinctive new species. Recognizing then this should in fact be a new genus, he proposed the name Miltonia to it as a homage to Lord Fitzwilliam Milton, an English orchid enthusiast. Lindley states then that the limits between a number of Oncidiinae genera, Cyrtochilum, Oncidium, Odondoglossum, Brassia and Miltonia, at that time classified as Vandaea, were yet to be perfectly established; although closely related, the differences should possibly be: Oncidium has a column with two ears and labellum distinctively lobed; Miltonia has a column with two ears and an entire labellum partially united to the column base; Odontoglossum and Cyrtochilum have winged columns and entire labelli but the former has it partially united to the column; and Brassia does not have any appendages on the column. It is interesting to notice that despite Lindley described the genus Aspasia in 1833, which is the most closely related to Miltonia, both by flower and vegetative morphologies, he did not mention it on Miltonia description.[9]

Three other Botanists were working with Miltonia species around the time Lindley described this genus. All recognized these plants should be classified under a new genus and, as communications were slower then, all proposed new genera: Knowles and Westcott also received also a plant of M. spectabilis and, just one month after Lindley, proposed for it the genus Macrochilus, calling the species Macrochilus fryanus;[10] the other one was Rafinescque who, in 1838, decided that the Oncidium russellianum already described by Lindley in 1836 should be under another genus and created for it the genus Gynizodon.[11] Both Macrochilus and Gynizodon are synonyms of Miltonia and no other species has ever been submitted to them.[2]

(CC) Photo: Dalton Holland Baptista
Miltonia russelliana
This is the less showy of Miltonia species because its sepals and petals do not really open, being always bent over the column, revealing only the lighter tip of its purple labellum.
(CC) Photo: Dalton Holland Baptista
Miltonia clowesii
This species has the same color pattern of M. russelliana, however, with a whiter labellum. On the other hand, M. clowesii flowers pointed segments are larger and wide opened making it look like a spider.

As Miltonia species are common plants, comparatively large, with also large flowers of bright colors, that, moreover, are spread mostly over an area of early settlements in Brazil all species but one were already described in 1850; six of them by Lindley, M. regnellii by Reichenbach[12] and M. moreliana by Achille Richard.[13] Despite the early description of M. moreliana in 1848, and two other as M. rosea by Lemaire in 1867,[14] and as M. warneri by George Nicholson in 1886,[15] Arthur Henfrey reduced it to a variety of Miltonia spectabilis in 1851,[16] and as such it was considered until 2002, when Cássio van den Berg reestablished it as a distinct species.[17] The last Miltonia species to be discovered was M. kayasimae, found by an orchid collector not far from the city of São Paulo, in an area around nine hundred meters of altitude nearby the top of Serra do Mar mountains. It was named after their collector by Guido Pabst in 1976. So far very few plants were found, all living at the same area.[18]

Since the genus Miltonia was established, many species, now classified under a number other genera, were submitted to it. The most noticeable cases were four of the five species of Miltoniopsis,[19] a genus proposed in 1889 but only really accepted in 1976.[20] Despite its somewhat similar flowers, Miltoniopsis are from cooler forests on the Andean slopes closely related to Cyrtochilum and only remotely related to Miltonia. Also five of the six Miltonioides species were occasionally considered as Miltonia until 1983 when Brieger and Lückel proposed this genus for them.[21] These are species of more delicate and narrower flowers, from Mexico and Central America, which some taxonomists claim might be better classified under the genus Oncidium to whom they are closely related.[22] The last common species which was occasionally classified under Miltonia is Chamaeleorchis warszewiczii,[23] which is related to Oncidium and some taxonomists identify as Oncidium fuscatum.

In 1983, Brieger and Lueckel, considering that four species of Miltonia, M. candida, M. cuneata, M. kayasimae and M. russelliana, show the junction of the labellum with the column in a different angle than the other species, proposed the genus Anneliesia for them.[24] Although this four species form a small sister clade to the rest of Miltonia species, the difference did not seem important enough to justify the acceptance of this new genus, therefore this proposal has not been generally accepted by the scientific community.[2]

In 2001, based on molecular analysis, Norris Williams and Mark Chase, transfered a species previously classified under the genus Oncidium, as O. phymatochilum, to Miltonia.[25] As this species shows a morphology that closer to Oncidium species than to Miltonia, because of its small yellowish flowers and highly branched inflorescence, this result and following transfer was a great surprise to most taxonomists. In 2005, Eric Christenson suggested a new genus and the name Phymatochilum brasiliense for it.[26] There is no consensus about the name to be generally accepted as yet.

Molecular analysis show that Miltonia most closely related genus is Phymatochilum and then Aspasia, Brassia and Ada, which are the most important genera included in this that is one of the eight clades that form the subtribus Oncidiinae of tribus Cymbidieae.[22]

Species

(CC) Photo: Dalton Holland Baptista
Miltonia cuneata
This is the hardest to grow among all Miltonia species, however, being all other very easy, this is not really a hard one. This species is the one that needs cooler conditions.
(CC) Photo: Dalton Holland Baptista
Miltonia kayasimae
This was the last species to be described and is the rarest one having being found just a couple of times in São Paulo State, in Brazil.

The species of Miltonia show many differences to each other and are very easy to identify, therefore, just the most evident differences are mentioned here; more details are given on individual species articles. The species are presented here according to their morphology and this order keeps no correspondence with phylogenetic relationships.

Regarding vegetative morphology Miltonia moreliana and Miltonia spectabilis can be immediately separated from the rest because their much flatter pseudobulbs, longer rhyzome and inflorescences completely covered by flattened bracts that bear only one highly flat flower. These are the species with largest flowers in the genus. They are closely related and usually are recognized because the flowers of M. moreliana usually have dark purple petals and sepals and the lip of a lighter bright purple while M. spectabilis has very light purple or white petals and sepals and a purple veined labellum, however, the real technical difference among the species is on the proportions of their segments which are much wider. Despite colors are often mentioned to identify species they are not accepted by taxonomy as enough to establish distinct species by themselves.[17] All other Miltonia species have similar vegetative appearance and only can be positively identified by their flowers.

Three species are very unique: Miltonia flavescens has the most narrow flowers, almost star shaped, with all segments of straw color with some purple bots on the base of petals and sepals which are more intense on the labellum almost forming stripes;[1] Miltonia candida is the only species with a labellum that embraces the column in a way that reminds the Cattleya species;[27] Miltonia russelliana is the less showy of Miltonia species because its sepals and petals do no really open, being always bent over the column, revealing only the lighter tip of its purple labellum.[8]

Miltonia regnellii shows the widest flower color variation among all Miltonia species; they can vary from white to yellow, pink and lilac with labelli also varying from white to dark purple. The flowers open in succession and slightly resemble the ones of M. spectabilis although much smaller. They actually are the Miltonia species with the smallest flowers.[12]

Miltonia kayasimae and Miltonia cuneata are somewhat similar and possibly are closely related, both have straw color petals and sepals almost entirely covered by large brown stains and white labelli, however, they show different proportions on the flowers segments. M. kayasimae has much wider petals and sepals and smaller labellum which, moreover, has a larger and more salient and complex entirely purple callus on its base which is delicate, more straight and simple, and just occasionally purple dotted on its apex on M. cuneata.[18]

Miltonia clowesii has the same color pattern of M. russelliana with light yellow greenish brown sepals and petals completely covered with large darker dots or stains and labellum of bright purple at the base and lighter apex, however here they are whiter. On the other hand, M. clowesii flowers pointed segments are larger and wide opened making it to remind a spider.[28]

Natural hybrids

(CC) Photo: Dalton Holland Baptista
Miltonia × bluntii
This is the natural hybrid of M. spectabilis and M. clowesii.

Considering its limited number of species, it is surprising that eight natural hybrids of Miltonia are currently known, a number that almost equals the number of species and also implies that the most important pollinator of the majority of the species possibly is the same. As the crossing of two species uses to produce variable plants most of these hybrids have been described more than once and some have three or four synonyms. M. spectabilis is the species which has produced the largest number of hybrids, five: Miltonia × bluntii when crossed with M. clowesii, Miltonia × cogniauxiae with M. regnellii, Miltonia × flava with M. flavescens, Miltonia × leucoglossa with M. candida and Miltonia × rosina with M. cuneata,[2] furthermore it is possible there is also one with M. moreliana, which has not yet been described because M. moreliana itself was earlier considered a variety of M. spectabilis.

M. candida, besides the hybrid already mentioned with M. spectabilis, also produced two others: Miltonia × binotii with M. regnellii and Miltonia × lamarckeana with M. clowesii.[2] The remaining hybrid, Miltonia × peetersiana used to be considered a synonym of M. × bluntii but because M. moreliana is now a species distinct from M. spectabilis it is its hybrid with M. clowesii, which has entirely purple flowers instead the one with light brown petals and sepals.[29]

Culture

Despite being easy to grow Miltonia species tent to be subject to spots on their thin leaves generally caused by fungi proliferation and normally, when exposed to the amount of light they need to a full bloom, their foliage gets a little bit too yellow colored, although they should never be exposed to full sunlight. Finding the right balance of light exposure to avoid the yellow leaves but still produce nice blooming is important and with some tentatives the grower will succeed. They are not much sensitive to temperature but it varies according to their origin M. cuneata being the one that grows cooler and M. moreliana the warmer grower all under intermediate temperature with at least 10ºC of variation between day and night. Despite they show a rest period after blooming, Miltonia always need to be watered, more abundantly during active growth. they need at least 65% of humidity and good ventilation all the time. Moderate weekly fertilizing with a balanced formula is beneficial during active growth. They may be potted in a compost of half-chopped Sphagnum, peat, and some medium sized lumps of charcoal, or mounted on plaques of vegetable fiber, however if mounted they will need more frequent waterings.[30]

References

  1. 1.0 1.1 Johnson, Andrés E. (2001). Miltonia flavescens in Las orquídeas del Parque Nacional Iguazú, 130. Fundación Vida Silvestre Argentina, Buenos Aires. ISBN 9509725412
  2. 2.0 2.1 2.2 2.3 2.4 Govaerts, Rafaël et al: World Checklist of Orchidaceae. The Board of Trustees of the Royal Botanic Gardens, Kew. Published on Internet (Access in March 2009).
  3. 3.0 3.1 Miller, David; Richard Warren; Izabel Moura Miller & Helmut Seehawer (2006). Miltonia in Serra dos Órgãos sua história e suas orquídeas, 323-6. Rio de Janeiro.
  4. Pabst, Guido & Dungs, Fritz (1978). Orchidaceae Brasilienses 2: 196, Brucke-Verlag Kurt Schmersow, Hildesheim. ISBN 3871050106
  5. Cogniaux, Célestin Alfred (1903). Miltonia in Flora Brasiliensis 3-6: 267-286. K.F.P. von Martius Ed.. Published on Internet, in Latin.
  6. Hoehne, Frederico C. (1940). Introduction in Flora Brasílica, Vol 12-1: 37-8. Secretaria de Agricultura de São Paulo.
  7. Lindley, John (1834). Cyrtochilum flavescens in Edwards's Botanical Register 19: t. 1627. England.
  8. 8.0 8.1 Lindley, John (1836). Oncidium russellianum in Edwards's Botanical Register 22: t. 1830. England.
  9. Lindley, John (1837). Miltonia spectabilis in Edwards's Botanical Register 23: t. 1992. England.
  10. Knowles, George B. & Westcott, Frederic (1837). Macrochilus fryanus in Floral Cabinet 1: 93. England.
  11. Rafinescque, Constantine S. (1838). Gynizodon russelliana in Flora Telluriana 4: 40..
  12. 12.0 12.1 Reichenbach, Heinrich G. (1850). Miltonia regnellii in Linnaea 22:: 851.
  13. Richard, Achille (1848). Miltonia moreliana in A.Rich., Portef. Hortic. 2: 38.
  14. Lemaire, Charles (1867). Miltonia rosea in Ill. Hort. 14.: t. 524.
  15. Nicholson, George (1886). Miltonia warneri in Ill. Dict. Gard. 2: 369.
  16. Henfrey, Arthur (1851). Miltonia spectabilis var. moreliana in Gard. Mag. 3:: 41. London.
  17. 17.0 17.1 Carlini-Garcia, L. A..; C. van den Berg; P. S. Martins (2002). A morphometric analysis of floral characters in Miltonia spectabilis and M. spectabilis var. moreliana. Lindleyana 17(3): 122-129. Published on Internet.
  18. 18.0 18.1 Pabst, Guido F.J. (1976). Miltonia kayasimae in Bradea 2: 88.
  19. Godefroy-Lebeuf, Alexandre. (1889). Miltoniopsis in Orchidophile 9: 63. Argenteuil.
  20. Garay, Leslie A. & Dunsterville, Galfrid C.K. (1976). Miltoniopsis in Venezuelan Orchids Illustrated. 6: 278.
  21. Brieger, Friedrich G. & Lückel, Emil (1983). Miltonioides in Orchidee 34: 130. Hamburg.
  22. 22.0 22.1 Williams NH, Chase MW, Fulcher T, Whitten WM (2001). Molecular systematics of the Oncidiinae based on evidence from four DNA sequence regions: expanded circumscriptions of Cyrtochilum, Erycina, Otoglossum, and Trichocentrum and a new genus (Orchidaceae) in Lindleyana 16(2): 113-139.
  23. Senghas, Karlheinz & Lückel, Emil (1997). Chamaeleorchis in Schlechter Orchideen I/C(33-36): 2305.
  24. Brieger, Friedrich G. & Lückel, Emil (1983). Anneliesia in Orchidee 34: 129. Hamburg.
  25. Williams, Norris H. & Chase Mark W. (2001). Miltonia phymatochila in Lindleyana 16: 284.
  26. Christenson, Eric A. (2005). Phymatochilum brasiliense in Richardiana 5: 195.
  27. Lindley, John (1838). Miltonia candida in Edwards's Botanical Register 24 (Misc.): 25. England.
  28. Lindley, John (1840). Miltonia clowesii in Sertum Orchididacearum: t. 34. England.
  29. Reichenbach, Heinrich G. (1886). Miltonia peetersiana in Gardener Chronicles, n.s., 26: 326.
  30. Baker, Charles O & Baker, Margaret L. (2006), Miltonia in Orchid Species Culture Oncidium/Odontoglossum Alliance: 340-7, Timber Press. ISBN 9780881927757

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