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SUMMARY: The focus of this paper is three-fold. First, I shall examine the species of the genus Cycnoches from a historical and critical standpoint with a view to clarifying the status and validity of the various species. I make no pretense to be a trained botanist or taxonomist, but I hope through examination of the literature, personal communications with recognized experts and illustrations of the species, to present the latest information I could find bearing on this confusing genus and its fascinating species. Second, I shall examine the hybridization that has been done within the genus and with its related genera and discuss possible future directions. Third, I shall describe and comment on the awards that have been made to the species and hybrids of Cycnoches.

I have included a bibliography of sources and several appendices which I hope will be useful to anyone who is interested in further exploration of this genus and its relatives. There is a Classification Appendix A, which lists all of the species described in the genus Cycnoches with citation of taxonomic authority and place of original publication. This appendix also summarizes the conclusions of this paper about the currently valid species in the genus. As has happened in the past, further research may prove some of the species, now reduced to synonymy, to be valid species. New species almost certainly will be identified and described. At this point, I want to make clear that I have chosen to employ the older usage of Eucycnoches for the section containing the type species. The currently correct usage, according to the rules of nomenclature, is Cycnoches for this section, but in order to show that a reference is to the section rather than the genus, I have used Eucycnoches for the section throughout this paper.

NOTE: Sources are cited in the text of this paper by referencing the author and year of publication. These citations refer to the Bibliography except for references to publication of original type descriptions. These are found in the Classification appendix. This approach eliminates the need for lengthy references and footnotes in the body of the paper.

The genus Cycnoches was erected by John Lindley in 1832 based on a single flower which he received from the nurseries of Conrad Loddiges & Sons, who were, at the time, one of the leading orchid growers and importers in England. The plant was one of numerous orchids sent in 1830 to Loddiges by John Henry Lance, Esq., an English barrister or solicitor who was living in Surinam. I was not able to locate much information on Lance so he appears to be a somewhat mysterious personality whose presence in a Dutch territory seems a bit strange. He supplied a number of new species to Loddiges, before his return to England in 1834. Among them was Oncidium lanceanum, which Lindley named for him.

Lindley identified the flower from Loddiges as not only a new species, but a new genus. The genus was named Cycnoches, from the Greek words for swan and neck, alluding to the supposed similarity of the column to the neck of a swan. The species was named loddigesii for the firm who provided Lindley with the flower (Lindley 1832).

In August 1836, Lindley received a flower for identification from an English grower, Mr. Wilmer. Lindley identified it as a new species of Cycnoches because of differences in shape of the floral parts, and lack of fragrance ( C. loddigesii is very fragrant). He named it Cycnoches cucullata. Shortly afterward, in the autumn of 1836, at the garden of the Horticultural Society, a plant of Cycnoches flowered with two inflorescences. One bore the fragrant flowers of C. loddigesii and the other the scentless flowers of C. cucullata. All of these plants had originated from the shipment Lance sent to Loddiges' nurseries. This occurrence began the development of the idea that Cycnoches, along with Catasetum, was in a "sportive condition," and capable of producing different types of flowers at different times. It was another thirty years, before Charles Darwin (Darwin 1862 and 1874) would publish the identification and explanation of flowers of different sexes in these two genera. Cycnoches cucullata became the first of many Cycnoches species to be placed into synonymy with a previously described species. This really is somewhat disconcerting since at the time there was only one other species!

In 1837, another species of Cycnoches was described and figured by James Bateman as Cycnoches ventricosum. The plant originated from the vicinity of Ixtapa, Guatemala, and had been sent to him by George Ure Skinner, a British merchant, resident in Guatemala. The various orchids bearing the specific name skinneri and uro-skinneri are named for him. The plate of the plant (t. 5 Bateman 1842) shows an inflorescence now known to be male flowers and a seed capsule on the other side of the pseudobulb. Bateman remarked that the capsule had been attached to the plant when it arrived in England and described it as most interesting because of the huge size of the capsule and the minute size of the seeds. This capsule was eventually to play its part in the solution to another mysterious description in this same book (t. 40 Bateman 1842).

In 1838, a third species was described in Germany as Cycnoches chlorochilon. It had been sent from Caracas to Berlin. This same species also flowered shortly thereafter in Loddiges' nursery, and was pictured by Maund (Botanist, ii, t 54) under the name Cycnoches ventricosum. This plant, figured by Maund, was not Bateman's Cycnoches ventricosum, but the newly described Cycnoches chlorochilon on its first blooming in England. This is the first of many times that these two species would be confused.

In 1840, a fourth species of Cycnoches was published by Lindley under the name of C. maculatum. This is the first species published of what would be the section Heteranthae. It was discovered in Venezuela by one of the collectors of Hugh Low & Co., another famous orchid nursery of the time and a vigorous competitor of Loddiges. So far, except for the misidentified flower by Maund, and the quickly identified error of Cycnoches cucullata, the situation within the genus was not very confused. This was shortly to change and real confusion would set in. The years 1842 and 1843 would shake Lindley's and other taxonomists' world.

In July 1842, Bateman published another new species of Cycnoches (t. 40, Bateman 1842). John Lindley (1843), Robert Allen Rolfe (1909d) and others quote verbatim the history of this species as described by Bateman. It probably is one of the most fascinating stories, not just in the history of orchids, but in botany as a whole. Bateman stated that among Skinner's earliest collections of orchids in Guatemala was a plant which is described as joining the long, pendulous inflorescences of Gongora with the growth habit of Cycnoches. Bateman quickly asked Skinner to send some of the plants to him. The plants arrived and when they bloomed in England, the flowers were identified as the already known Cycnoches ventricosum. A mix-up was assumed and Skinner was asked to send plants a second time. When these bloomed, again they were identified as Cycnoches ventricosum. Skinner was asked a third time to send the new species and since he was returning to England on a visit, he took the plants with him under his personal supervision. On the voyage home, a plant bloomed and the flowers were indeed the new species. Skinner was sure that this time there would be no confusion. The flowers were exhibited to Bateman on arrival and the plant was put into Bateman's plant house or "stove" as it was called then. The new species was described as Cycnoches egertonianum (Bateman 1842, t. 40). The plant grew well and when it came time to flower, it produced the, by this time, well-known flowers which were identified as Cycnoches ventricosum. Bateman stated that this inflorescence was still hanging on the plant, when a second inflorescence was produced and it was the much sought-after Cycnoches egertonianum, which of course was radically different from the flowers first produced. No explanation could be offered to explain this dual flowering at that time. Cycnoches ventricosum was known to produce seed in its own right as was illustrated by the first plant of C. ventricosum in t. 5 of The Orchids of Mexico and Guatemala (see above) which had arrived in England with a seed pod attached. The new plant is shown in t. 40 in Bateman 1842, and clearly displays the long male inflorescence of Cycnoches egertonianum on one side and male flowers of Cycnoches ventricosum on the other. This presented an inexplicable mystery, especially since Bateman personally vouched for the authenticity of the plate. The mystery took fifty years to solve, and we shall discuss the resolution later in the paper.

The following year, John Lindley (1843) discussed this plant in the Botanical Register. He quotes Bateman's story as described above and says that many persons felt convinced some mistake had been made in this plate. Then, Lindley asked "But what is impossible in nature?". He says that in September of 1842, he received an inflorescence which bore flowers of Cycnoches ventricosum and Cycnoches egertonianum intermixed (his emphasis). A woodcut of the inflorescence is included (shown at left on page 5).

Lindley states the plant was exhibited at a meeting of the Horticultural Society, noting that it had been purchased as Cycnoches ventricosum. The inflorescence bears five flowers and I quote Lindley's description of the flowers. "Here it will be seen that fig. 2. is nearly Cycnoches ventricosum, but its lip is here and there raised into warts, which are the beginning of the lobes of Cycnoches Egertonianum, and moreover some of the dark purple of the latter is appearing at the base of the column and the tips of the sepals. At fig. 3. the purple of Egertonianum is displacing the green of v entricosum, the sepals are rolling back, and the labellum is almost wholly changed, but the sepals are still those of C. ventricosum. At Fig. 4. and 5., the transformation is complete. Another curious point in this instance is that the transformations occur in no certain order. The lower flower on the spike, No. 1, is more Egertonianum than ventricosum; that which succeeds, No. 3, is more ventricosum than Egertonianum; and 4 and 5, the last on the spike, are wholly Egertonianum. What with such cases as this, ...all ideas of species and stability of structure in the vegetable kingdom, are shaken to their foundation" (my emphasis). This inflorescence seemed to confirm the idea that Cycnoches species were a group in a "sportive condition" and might at any time produce flowers of different sorts from the same plant.

The extent of Lindley's frustration and doubts about Cycnoches caused by these two plants is illustrated in his description of Cycnoches egertonianum var. viride (Lindley 1846). I am going to quote this almost completely, since it illustrates so well the confusion that this genus was causing the leading taxonomic authority of his day and others. Some of this confusion exists to this day even after the many clarifications that have been published, as we shall see later in this paper. After the description of the variety, Lindley goes on to ask "But what is C. Egertonianum itself?" He briefly describes the dual-flowered plant illustrated by Bateman and concludes " C. Egertonianum is then a 'sport,' as gardeners say, of C. ventricosum." Lindley's next question is

But what again is C. ventricosum? Who knows that is not another 'sport' of C. Loddigesii, which has indeed been caught in the very act of shewing a false face, something wonderfully suspicious, all things considered, and justifying the idea that it is itself a mere Janus, whose face is green and short on one side, and spotted and long on the other. Then, if such apparently honest species as C. Egertonianum, ventricosum, and Loddigesii are but counterfeits, what warrant have we for regarding the other so-called species as not being further examples of plants masquerading with false faces? For ourselves we cannot answer the question; nor should we be astonished at finding some day a Cycnoches no longer a Cycnoches, but something else; perhaps a Catasetum. If one could accept the doctrine of the author of the 'Vestiges,' it might be said that in this place we have found plants actually undergoing the changes which he assumes to be in progress throughout nature, and that they are thus subject to the most startling conditions only because their new forms have not yet acquired stability. Be that as it may, it will be as well to state in this place what the so-called species are of this extraordinary genus, and what the changes are which they have been seen to undergo.

1. C. Loddigesii... Sports by producing smaller broad-lipped flowers without scent, and with a very short club-shaped column.

3. C. chlorochilon...Has not been observed to sport: but is probably a mere variety of C. ventricosum.

I find this article one of the most amazing publications I have seen in the literature on orchids. At this point, the genus is only twelve years old and the doubts about it are extensive. As I said above, the explanation of the dual, male-flowered plant illustrated by Bateman (1842, t. 40) will be given later in its chronological place, but I do at this point want to discuss the second inflorescence with the five flowers of different types (Lindley 1843). Paul Allen (1952) in his revision of the genus Cycnoches explains that the inflorescence was the first to be seen of Cycnoches egertonianum with several male flowers and one of the rare hermaphroditic form produced together on one spike. I have examined this woodcut in its original publication (see page 5 above) and Lindley's description quoted. I wonder about the two flowers numbered 1 and 3. It is now obvious that flower number 2 is indeed the hermaphroditic or "perfect" form of Cyc. egertonianum and numbers 4 and 5 are normal staminate flowers. However, the form of the other two is not typical of the staminate Cyc. egertonianum. The sepals and petals are much broader and flatter than typical. The columns, especially of number 3, appear to be thicker and shorter than the typical staminate forms. If we assume that the drawing is accurate, it would seem that perhaps flower numbers 1 and 3 might be a different example of the so-called hermaphroditic form. As we shall see later, there is indeed more than one type of "perfect" flowers in the genus and I have the feeling that numbers 1 and 3 may be of the intermediate type described by Allen ( op. cit.) which is closer to the normal staminate flowers. This type is even rarer than the usual hermaphroditic form which is closer in appearance to the female flowers except for the additional "bumps" on the lip, some of the color usually found on the staminate form and, of course, the presence of both pollinia and a stigma. This would make the inflorescence even more amazing than Allen thought originally.

Despite the confusion and doubt expressed by Lindley about the genus and its species, to paraphrase an old saying "you can't keep a good taxonomist down" and so publication and description of new species in the genus continued, even though there were suspicions about the status of Cycnoches and its "sportive condition". The next species to be examined illustrates how deeply the suspicions of the genus were and how these could spread confusion to our day and to the A.O.S. Awards system.

Between 1846 and 1852, six more species were described to Cycnoches (three of which are now referred to other genera), but it is Cycnoches warscewiczii which I want to examine now. In 1852, Heinrich Gustav Reichenbach, normally referred to in botanical literature as Reichenbach filius because his father was also a prominent taxonomist, was twenty-four years younger than Lindley and was just beginning what was to be a long and illustrious career in orchidology (Reinekka 1972). I find it interesting that 1852 coincidentally marks the publication of the first two species of Cycnoches described by Reichenbach f. and the last published by Lindley. Although Lindley lived until 1865, illnesses in 1851 and 1862 had their effect on his normally prodigious amount of work. He continued to write and publish until his second illness in 1862. Lindley and Reichenbach f. had been friends and colleagues since 1849 and according to Reichenbach's account, he was the last person except Lindley's family and doctor to see Lindley before his death on November 1, 1865 (Reinekka 1972).

One of the species published by Reichenbach in 1852 was named Cycnoches warscewiczii in honor of the famous orchid collector Josef Ritter von Rawicz Warszewicz who had sent the plant to Reichenbach. Von Warszewicz is memorialized in the specific epithet warscewiczii in a number of other species which he collected and sent back to Europe. Cycnoches warscewiczii was described from a single pistillate flower which Warszewicz had collected in Chiriqui Province in Western Panama, the part nearest the Costa Rican border. At that time, Panama was a province of Colombia. In this account I am following Paul Allen (1952). Reichenbach was evidently influenced by the then current theory that Cycnoches was in a "sportive condition". He closes his description with the comment: "I wonder if (this) is not one of the wonderful dimorphic forms that haunt this genus." In keeping with this idea, as late as 1866 (Beih. Orch. Central Amer. p 23), he listed the species as (sic) " Cycnoches--sexus Cycnoches ventricosi Bat.?-- Warscewiczii Rchb. f.". As he had specified in his will, Reichenbach's herbarium was reopened twenty-five years after his death. The herbarium contains a drawing of what is probably the type. It shows a single pistillate flower and the column of a staminate flower, both of the section Eucycnoches, labeled "Chiriqui, Cordillere. Leg. Warscewiczii."

As if description from a single female flower was not confusing enough, there is another drawing in the Reichenbach herbarium of two or more other collections of several very different, smaller, pistillate flowers typical of the section Heteranthae. They also bear the inscription " Cyc. Warscewiczii." This demonstrates that Reichenbach was a believer in the "sportive condition" of the genus and its ability to produce flowers of almost any type, at any time.

Rolfe (1909d) recounts that in 1879, The Gardeners' Chronicle published a plate showing a female flower on one side of a plant of Cycnoches, and a section of an inflorescence on the other side with typical Heteranthae male flowers. The plate (12:493, fig. 78) is based on a plant exhibited at a meeting of the Royal Horticultural Society on October 14 by a Mr. Bull. The plant was identified as Cycnoches warscewiczii Rchb. f. and Mr. Bull received a Botanical Certificate, R.H.S. for the plant. Rolfe notes that the plant is not the C. warscewiczii of Reichenbach. I believe that this plant and the plate may be the origin of the misidentification of the plant which was given two A.O.S. awards as " Cycnoches warscewiczii var. warscewiczii" 'Jan's Swan'. The awards will be discussed in Part III under the A.O.S. awards. We shall see below, in the discussion of the articles by Rolfe (1909d) and Schlechter (1916) which organized and described the species of the Cycnoches, that doubt about the species was still in the mind of the next generation of taxonomists. As Allen (1952) said "error breeds error." The validity and correct assignment of Cycnoches warscewiczii were to wait until the 1940's when the name was revived by the Ames Herbarium in Cambridge and the identity of Cycnoches warscewiczii was confirmed by comparison with the types in Reichenbach's herbarium. Reichenbach willed his herbarium to the Naturhistorisches Museum in Vienna after his death in 1889. Since Reichenbach's will specified that his herbarium was to be sealed for twenty-five years after his death, neither Rolfe nor Schlechter had access to the material. This sealing of Reichenbach's work was mentioned several times by Rolfe as a great hindrance to taxonomic work. It evidently was a source of great irritation to Rolfe. It has been suggested that Reichenbach decreed the sealing of his herbarium because of jealousy or personal animosity toward Rolfe. Rolfe was a self-taught expert on the orchids, having begun his career as an assistant gardener at Kew, who through industry, hard work and close observation, rose to scientific eminence (Reinekka 1972).

A major step toward the clarification of some of the problems in the genus Cycnoches and its wonderful dimorphic (actually tri- or polymorphic) flowers, occurred in 1862 with the publication by Charles Darwin of the results of his research about different sexual forms of Catasetum tridentatum (Darwin 1862) and of his now classic work On the various contrivances by which British and foreign orchids are fertilised by insects, as it was originally titled. I have used the expanded and updated version that he published in 1877 under the title The various contrivances by which orchids are fertilised by insects which was reprinted with a useful foreword by the University of Chicago Press in 1984. Darwin's work was the first serious investigation of the pollination of orchids. Darwin was fascinated and intrigued by the mechanisms of orchids to avoid self-pollination. His studies and conclusions on European orchids and their pollination are in general still valid. His special interest however, was in the complex pollination mechanisms of tropical orchids. All of his work was done in the laboratory with a relatively small number of specimens he was able to obtain from European sources. He was dependent on observers who lived in the tropics and who, unfortunately, supplied him with faulty information in some critical areas. Additionally, he worked without knowledge of the pollinators -- the insects who were performing the task of transporting the pollen to the stigma. Because of these factors, Darwin made some errors, but, despite errors, his work was a landmark step forward. The idea that there were male and female flowers in the genera Cycnoches and Catasetum had been noted previously as we saw above in Reichenbach's notation about Cycnoches warscewiczii. It was Darwin's work, however, which placed this idea on a factual basis with scientific proof.

Darwin's work in the Catasetinae was concentrated on species of Catesetum, but he recognized the relationship within the Catasetinae and discussed the genera Mormodes and Cycnoches in Chapter VII (1874). Although I admit to a prejudice or bias, I especially enjoy Darwin's first sentence of Chapter VII, which I quote. "I have reserved for separate description one sub-family of the Vandeae, namely the Catasetidae" (as it was called at that time), "which must, I think, be considered as the most remarkable of all Orchids." At that time species we now know as Catasetum had been placed in three different genera, Monachanthus, Myanthus and Catasetum. Their situation was as confused, if not more so, than the status of the Cycnoches. I will not recount the details of Darwin's research, but in the course of his work on the pollination mechanism in these three genera, he reached the conclusion that the flowers of the five Catasetum species he had received represented male forms without functional female parts. He received preserved specimens of Monachanthus viridis and Myanthus barbatum for his research . He was also working with field reports from Trinidad from two sources who were considered reliable. They had reported that Catasetum tridentatum, Monachanthus viridis and Myanthus barbatum grew intermixed on the same plants. After his examination of Monachanthus viridis and Myanthus barbatum, Darwin concluded that Monachanthus viridis was the female flower and that Myanthus barbatum was the hermaphroditic form of Catasetum tridentatum. The latter conclusion was wrong, but it was based on inaccurate field information which led Darwin to incorrect conclusions. He was correct that the flowers of Monachanthus viridis he was working with were female. As proved by Rolfe (1889), however, his conclusion about Myanthus barbatum was wrong and it was not a hermaphroditic form. Catasetum tridentatum is now identified as synonymous with Catasetum macrocarpum (Romero and Jenny, 1993) which belongs to the section of Catasetum whose flowers are non-resupinate with a "helmet"-shaped lip and therefore bear a resemblance to the female flowers of all the Catasetum species. This section could be described as the Catasetum equivalent of the Cycnoches section Eucycnoches in that within each section the female flowers are similar to the male flowers. The idea that there were three different sexual forms of flowers in the Catasetum species has proved to be correct, but Darwin's information led him to misidentify the one with which he worked. Rolfe in his paper on Darwin (Rolfe 1889), as noted above, proved that Myanthus barbatum (now called Catasetum barbatum) was not hermaphroditic, but was a male flower. Rolfe reached the erroneous conclusion that a hermaphroditic form did not exist, only male and female. This is incorrect as we now know, so in the end Darwin's idea was correct, even if the information he based it on was wrong.

In the final short section of the chapter, Darwin wrote on Cycnoches ventricosum. He was never provided with female flowers and so worked only from male flowers. His conclusions after examining them were that they were indeed male flowers, but he said it might be possible that after removal of the pollinia and the membrane which covered the stigmatic surface (which is vestigial in the male flower), the flower might be pollinated as was the case with Mormodes. He made reference to Bateman's and Lindley's observations and the reports of the "sporting" nature of Cycnoches ventricosum and C. egertonianum. Darwin then concluded that, based on the analogous differences in Catasetum, "we are tempted to believe that we here have male, female, and hermaphrodite forms of the same species of Cycnoches." Again, because of misleading information and lack of material, Darwin's conclusions were incorrect. There are three sexual forms of Cycnoches, as postulated, but they are not the three separate species about which Darwin was given information.

Although, I shall not go into detail here, in the period after 1852 until the publication of Rolfe's Revision of the genus Cycnoches (1909d), ten more species were described in the genus. In 1909, R.A. Rolfe published the first comprehensive organization of Cycnoches. Rolfe, in his Revision , was at last able to confirm the suspected error in Bateman's famous plate in t. 40 of Cycnoches egertonianum (Bateman 1842). The identification of female flowers of a number of species is another of the useful steps toward clarifying and reducing some of the confusion about Cycnoches which Rolfe incorporated into this paper (Rolfe 1909d).

Rolfe began with a review of the history of the genus. I have already covered the material through Darwin's proof of the sexuality of flowers in the Catasetinae and the description of the award in 1879 to a so-called Cycnoches warscewiczii. Rolfe recounted the appearance and identification of female flowers of Cycnoches pentadactylon in 1889 and, in 1891, of the female flowers of Cycnoches chlorochilon. In that year, the female flowers of C. chlorochilon appeared and were confirmed in two different collections. They appeared twice in a Belgian collector's greenhouses on plants imported from Venezuela. Three female flowers also appeared in an importation of Sanders' to England. The actual confirmation of the identity of both male and female flowers of C. chlorochilon was their appearance on the same plant in Belgium. Their identity was now beyond doubt and confirmed the identity of the earlier flowering in Belgium and the three at Sanders'. In 1891, Rolfe also described a new species based on a female flower sent to Kew for identification and in the same season, the subsequent production of male flowers on the same plant. This was the first species to be described initially from both male and female flowers. The new species was named Cycnoches rossianum in honor of Mr. H.J. Ross who had purchased the plant under the name Cycnoches warscewiczii (!!). (This species has now been reduced to synonymy with C. egertonianum.)

At this point in his paper, Rolfe stated that it had become increasingly evident that some mistake had been made in Bateman's original description and figuring of Cycnoches egertonianum (t. 40 in Bateman 1842, the famous plant with two different male inflorescences). Rolfe said that he examined all the materials available up to the time he was writing, including his own description of Darwin's mistake (Rolfe 1889). Then, after discussion of Bateman's story, Rolfe came to the conclusion that the flowers that were identified as Cycnoches ventricosum on the plant could not have been drawn from life. He concluded that the so-called Cycnoches ventricosum flowers would have withered by the time the inflorescence of the true male flowers of Cycnoches egertonianum appeared. Therefore, Miss Drake, who did the drawing, used the fresh flowers of a male Cycnoches ventricosum from another plant to complete the drawing. This would not have seemed strange at the time, since the flowers had originally been identified as C. ventricosum. Rolfe published these conclusions in the Gardeners' Chronicle (1892, xi. pp. 204, 205) and then it occurred to him to write Bateman who was still alive at that time. Bateman replied that although he remembered the circumstance perfectly, he could not remember exactly how the drawing was made, but he was inclined to agree to the suggested explanation.

Rolfe recounted that three years later, his conclusion was confirmed. In the autumn of 1894 a living plant of Cycnoches was received at Kew from Costa Rica. In the spring of 1895, this plant produced a new growth from the top of the old pseudobulb. The new plant was removed and grown side by side with the old pseudobulb. In April 1895, one of them produced a long raceme of purple flowers identical with C. egertonianum. In June 1895, the second plant produced two green flowers which were clearly female. The paper includes drawings of the female flower which Rolfe now had definitely identified as C. egertonianum, along with the male flower and, for comparison, the male and female flowers of Cycnoches ventricosum. He states that "a glance will show how distinct it" (the female flower of C. egertonianum) "is from the male of C. ventricosum although the flowers are green in both". Plates III and XI to this paper illustrate these two species. There are clear differences, not only in the column structure, but also in the fact that the female flower of C. egertonianum does not show the strongly reflexed sepals and petals characteristic of both the male and female flowers of C. ventricosum. Thus, fifty-three years after the publication of the original description, the solution was published.

The final part of Rolfe's paper is devoted to the description and naming of the two sections of Cycnoches, the Eucycnoches and the Heteranthae and the placement of the sixteen species considered valid at that time into these sections. Within the sections, the species are divided into two groups. Synonyms, as they were considered at that time, are also identified. The first group in each section is the species with both male and female forms known. The second group is the species with only male forms known. In conclusion, two female flowers were listed whose identities were in doubt at the time, with a possible identification of the species to which they might belong. Botanical citations and places of publication are given, including all the plates showing the species that had been illustrated up to his time. He placed six species in the section Eucycnoches, three of which, C. loddigesii, C. ventricosum and C. chlorochilon, had both sexes identified; three of which, C. lehmannii, C. haagei (sic) and C. versicolor, had only the male flowers identified at the time. I note at this point that C. versicolor is now considered to be synonymous with C. haagii. In the section Heteranthae Rolfe included ten species, six of which he identified as having both male and female flowers known and four whose female forms were not then identified. The six species for which he identified both male and female flowers were C. egertonianum, C. pentadactylon, C. stelliferum, C. aureum, C. rossianum and C. densiflorum. The four species he listed with only male forms known were C. maculatum, C. dianae, C. glanduliferum and C. peruvianum. The two unknown female flowers at the end of Rolfe's description were tentatively assigned to C. peruvianum and C. maculatum.

Because of the confusion which at that time surrounded Cycnoches warscewiczii, I wish to include Rolfe's discussion of this species, which you will have noticed is not included as a separate species in the Revision. Rolfe actually listed Cycnoches warscewiczii twice, once as C. warscewiczii Hort., giving as its citation the plant awarded a Botanical Certificate by the R.H.S. in October, 1879, and which was published in The Gardeners' Chronicle as cited above and five other places by Rolfe. He noted that these flowers were quite distinct from the C. warscewiczii of Reichenbach and stated his belief that Cycnoches warscewiczii Rchb. f was the female form of Cycnoches aureum. Rolfe said an authentic flower of C. warscewiczii was preserved in Lindley's herbarium and that it was almost twice as large as the female flowers of the C. warscewiczii Hort. He noted that the flower described by Reichenbach as Cycnoches warscewiczii was collected by Warszewicz in Chiriqui province in Panama. He said that the Lindley herbarium contained part of an inflorescence with two male flowers labeled by Reichenbach " C. aureum, Chiriqui Vulc. 6-8000 feet, Oct., Warszewicz". From these facts, Rolfe inferred C. warscewiczii Rchb. f. to be the female form of C. aureum. He noted that Reichenbach himself suggested that it might be the female flower of C. ventricosum. While Rolfe's deduction was logical, as we now know, it was incorrect -- and so was Reichenbach's suggestion. Rolfe, of course, knew the true female form of C. ventricosum, as described earlier in his paper, and so he knew Reichenbach was wrong.

I have gone into some detail about Rolfe's Revision, but since it is the first attempt to outline and organize the genus, it is a major landmark in any study of Cycnoches. While Reichenbach had earlier recognized that there were two logical divisions of the genus, it is Rolfe in this paper who established and named the two sections. He also summarizes well the knowledge of Cycnoches species at that time. Rolfe described one more species after publication of his Revision, C. cooperi which was published in 1913.

The next in the series of famous botanists and taxonomists to become interested in the genus Cycnoches was Friedrich Richard Rudolf Schlechter. In 1916, he published his paper Die Gattung Cycnoches Ldl . (The Genus Cycnoches Lindl.). As far as I was able to determine, there was no published translation of this paper into English when I began my research for this paper. Since Schlechter's paper represents the second publication of the genus as a whole, I felt the need to know what Schlechter said directly, without relying on later authors' summaries of what he said. I am fortunate to have a friend, Dr. Ursula Hoffmann of Lehman College - City University of New York, who was generous enough to translate the paper for me (Schlechter 1916). I wish to acknowledge at this point my gratitude to her for her translation and her friendship.

Schlechter began by noting that species of Cycnoches had begun to appear in German collections over the past few years and that their identification was quite difficult even for experts. Referring to the publication of Rolfe's Revision in 1909, Schlechter said that since many readers of Orchis either did not know or could not obtain the Kew Bulletin in which the article appeared, he would briefly summarize the history of the genus as presented by Rolfe. The sections of this paper, following the historical introduction, present a well-organized and detailed description of the female flowers of the two sections and their basic similarities, followed by an equally well-done summary of the male flowers of the two sections with emphasis on the differences in form between the sections. The genus' plant habit, habitat, culture and relationship to the other genera of the Catasetinae are then discussed. As a side note, in 1915, Schlechter had published the subtribe Catasetinae more or less as presently constituted, by removing what we now call the Cyrtopodiinae to a separate group.

The final part is devoted to a detailed key and description of all the species recognized as valid at the time of publication. It presents the sixteen species listed by Rolfe (1909d) plus Cycnoches cooperi which had been described by Rolfe in 1913, a total of seventeen species. Schlechter's revision is an important contribution to the knowledge of the genus Cycnoches. It represents the first key with consolidated detailed descriptions of the plant habit, geographical distribution, floral structure of the male flowers, notation of closely related species and their distinguishing differences. There are notes on the discovery and appearance in European collections of the various species. I do want to mention that, like Rolfe, Schlechter passes over C. warscewiczii Rchb. f., as a valid species because of its then doubtful status that originated with Reichenbach himself.o The synonyms listed agree with those of Rolfe. Schlechter's key along with Allen's (1952) form the basis of the key I have been working on. It is based on observable features of the male flowers. The part of the key for the section Eucycnoches is found at the end of this paper starting on page 66 below. I am still working on the key for the more complex and numerous Heteranthae.

According to Oakes Ames (1944), C. W. Powell who lived in the Panama Canal Zone in the early 1900's, was deeply interested in the orchids of Panama. In his efforts to identify them, he sent a number of specimens to R.A. Rolfe at Kew. Ames said that Rolfe was hesitant about supplying names because many of the species had been described by Reichenbach and, as already mentioned, Rolfe did not have access to Reichenbach's herbarium which was still sealed in Vienna. Ames said that Powell then turned to Schlechter for help with identification. From these plants, Schlechter identified eight more species in the genus Cycnoches: C. powellii, C. stenodactylon, C. guttulatum, C. pachydactylon, C. pauciflorum, C. tonduzii, C. amparoarum and C. brachydactylon. They were all published in Fedde's Repertorium in 1922, 1923 and 1924, but the work on the specimens had begun before World War I. With Schlechter's death in 1925, interest in Cycnoches seems to have declined. Cycnoches buchtieni was described by Kraenzlin in 1928 and C. espiritosantense Brade ex Hoehne was published in 1942 in Brazil. These were the last new species to be described until 1989. Cycnoches buchtieni was rather quickly transferred to the genus Polycycnis.

The only other work I wish to note on Cycnoches in this period, is the resolution to the problem of the status of Cycnoches warscewiczii. In the early 1940's, Oakes Ames of the Harvard University Herbarium identified Cycnoches warscewiczii, as the most widely-occurring Cycnoches of section Eucycnoches in Costa Rica and Panama. The male form of this species had been described and published by Schlechter (see above) as Cycnoches tonduzii based on material supplied to him by C.W. Powell in 1913. Photographs and drawings of material from Costa Rica and Panama of the male C. tonduzii and its female form were sent to Vienna to be compared with Reichenbach's herbarium material. Their identity as C. warscewiczii Rchb. f. was confirmed. Thus, this species was validated and placed in the correct section Eucycnoches (Allen 1952). Another problem was resolved although it took almost ninety years from the publication of the original description.

The event which was to have the most impact on our current understanding of Cycnoches came in the late 1940's and early 1950's with the resumption of the study of the pollination mechanism and the pollinators of the tropical American orchids by Paul H. Allen. Allen's early training was at the Missouri Botanical Gardens. Almost no work had been done in this field since Darwin's time (Darwin, 1862; 1874). Allen did work on the pollinators of several genera, but since this paper is concerned with Cycnoches, I shall not include any details about the work with other genera. With the assistance of Harry Dunn who lived in the Panama Canal Zone, Allen, who worked in Costa Rica, Panama and Honduras, devoted a lot of field work to Cycnoches. This research also led to the publication of a revision of the genus (Allen 1952). The paper presented details of the mechanics of pollination and the role that fragrance played in the process. He observed that the male bees were attracted to the fleshy callus at the base of the lip in Cycnoches warscewiczii. Following Darwin, who had based his information on the field work of Cruger, Allen reported that bees of the genus Centris were seen to gnaw on the fleshy callus at the base of the lip of Cycnoches warscewiczii. In the process of departing from the flower, the segments of the bee, posterior to the thorax, came into contact with the rostellar strap of the column. This caused the pollinia to be ejected and attached to the bee's posterior by the sticky disk at the end of the rostellar strap. Allen also observed that it took some 15 to 20 minutes for the strap to dry and straighten. This action placed the pollen in position to be caught by the notches on the column of the female flower. Further, he observed that it took some 30 minutes for the anther cap to dry sufficiently to be removed so that the pollen was finally exposed. He deduced that this delay was designed to insure ample time for the bee to visit a female flower. Allen also identified the function of the two deep notches which appear on the sides of the tip of the column of the female flowers. These notches, together with the broader, shorter shape of the column are the easiest characteristics used to identify female flowers from male flowers in the genus Cycnoches. The notches serve to catch the rostellar strap and, as the bee departs, strip the pollinia depositing them on the stigma.

Allen's paper offers the first detailed information I found published on the hermaphroditic forms of Cycnoches. He referred to the woodcut published by Lindley (1843) and the hermaphroditic flower number 2 shown (discussion of his analysis is found on page 5 and 6 above). Allen found that although the forms were sometimes mentioned in the literature, they were excessively rare in herbarium collections. He recorded a single specimen in the Ames Orchid Herbarium which came from Honduras and observed that it very nearly matched the flower in the Lindley plate (1843). He reported that another example appeared in Harry Dunn's collection in 1949. Allen identified all three of these flowers as Cycnoches egertonianum. Allen reported that in the fall of 1950, several hermaphroditic forms of Cycnoches warscewiczii appeared in his collection in Palmar, Costa Rica. Two of the hermaphroditic forms appeared on an inflorescence with two normal male flowers and two others appeared on a separate inflorescence. He observed that the two hermaphroditic flowers on the inflorescence with the male flowers had columns which were intermediate in form between the normal male and female flowers; however, they lacked the "critical notches" found on female flowers. On the separate inflorescence, the flowers had columns that were short and thick and completely typical of the female flowers, including the notches, except for the presence of an anther and pollinia. This is the first published report I found of two different forms of "perfect" flowers.

Allen said that the reason hermaphroditic flowers are so rarely found in herbaria may be at least partially explained by the fact that in both of his plants, the "abnormal" anther and pollinia were spontaneously ejected within about two hours after the flowers opened. This is very unlike normal male flowers which do not lose their pollinia for days, unless removed by a pollinator. Allen also observed that when normal male flowers lose their pollinia, they quickly wilt. The hermaphroditic flowers however remain fresh for many days afterward. Assuming that this is probably typical, Allen concluded the rapid loss of the pollinia is the reason so few hermaphroditic forms have been observed. With the pollinia missing, it would be easy to assume they are female flowers. Hermaphroditic forms of both C. egertonianum and C. warscewiczii are pictured in Allen's Revision, figures 17 and 18. Allen stated that it is probably highly significant that the perfect forms of each section should bear a striking resemblance to each other, and in fact, they are about what one would expect for two closely allied species in "one of the more conservative genera". He postulated that these may in fact be reversions to an ancient type and that they represent an approximation of the primitive condition of the genus before it embarked on the present "unique and divergent course", as Allen calls it. I have gone into some detail on these points because it is the first time this information, which appears to agree with current knowledge of the genus, was published.

The rest of Allen's paper (three of the five parts) is a thorough revision of the genus. The revisions of Rolfe (1909d) and Schlechter (1916) are more in the nature of logical and coherent organizations of the knowledge available at the time of their publication, rather than revisions such as Allen published. Rolfe and Schlechter made excellent and lasting contributions to the knowledge of the genus. In the light of later information which has been published since Allen's study, they have been justified in many cases (not all) just as Darwin (1862; 1874) was eventually proved to be correct in his conclusion of three sexual forms even through he was not correct in the details which led to the conclusion. Allen's Revision is devoted to the establishment of what he considered to be the truly distinct species in a logical attempt to accentuate what he saw as the very apparent relationships within the genus. His aim was to clarify the genus by reducing to synonymy the species which he concluded were described based on natural variations "in minor details".

Allen provides an excellent key to the seven species, two of which have two varieties each ascribed to the them, for a total of eleven valid species and varieties. The remaining twenty-five previously described species are reduced to synonymy with one of the eleven species and varieties he validates. Allen's key and descriptions are detailed, clear and well-written. They are extremely useful today when dealing with the species and so-called varieties he found valid. He recognized, and so stated, that future research and information might lead to the re-establishment of some of the synonyms as valid species, or at least distinct forms or varieties. It should be noted at this point that Allen based his revision on the examination of available type specimens, descriptions and very extensive field work with the species, all of which are perfectly valid methods of botanical research. Although he began the study of the pollinators, he did not carry this information into the examination of the validity of the described species. Later study of the pollinators and fragrance analysis has contributed the very information which Allen predicted might appear to change his findings to our latest view of Cycnoches. Recent studies in cladistics and molecular botany may contribute yet more information to change the present view of the genus.

In the process of writing his revision, Allen concluded that the most taxonomic confusion is found among species of the section Heteranthae. In the past, he stated, taxonomists "fixed on small variations" among the male flowers of the Heteranthae. These variations in the relative length of the basal claw of the lip, relative length and width of the apical lobe of the lip, and especially on variations in the length and shape of the finger-like marginal teeth of the lip, led to establishment of new species. He particularly criticized Schlechter for retaining as valid in his Revision what Allen considered to be doubtful species and adding to the situation in his description of the new species, seven of which are in the Heteranthae. None of Schlechter's new species survived in Allen's Revision. He also criticized Rolfe in addition to Schlechter for what he called the idea that any discernible deviation from past descriptions should be described as a new species. None of the species published by Rolfe survived Allen's Revision either. Chronologically, the latest species which survived was Cycnoches haagii, published in 1882 by the Brazilian Barbosa Rodrigues. Following is a summary of Allen's revision:

Eleven previously published species were placed in synonymy with Cycnoches egertonianum itself. Two more were placed in synonymy with C. egertonianum var. dianae, plus, of course, both C. aureum and C. dianae were reduced to the status of varieties of C. egertonianum. This is a total of fifteen species which Allen classified as variations of C. egertonianum. It should be noted that Allen added at the end of the description of Cycnoches egertonianum var. dianae that it was "probably a fairly good example of incipient speciation".

Beginning in 1959, Calaway H. Dodson and G.P. Frymire took up where Allen had left off in studies of the orchids of tropical America and their pollinators. Dodson and Frymire began their work in Ecuador, whereas Allen had done most of his work in Panama and Costa Rica (Dodson & Frymire 1961, Dodson 1962b). They quickly noted the key role that fragrance played in attracting pollinators. Since Cycnoches is one of the most fragrant of the genera, along with the other members of the Catasetinae, they did a lot of work with Cycnoches species, fortunately for the author of this paper. They observed that while the majority of plants, including orchids, attracted their pollinators with the offer of some type of food, many of the more advanced orchids developed various structures, odors or colors which imitated something important in the insect's life cycle. This was described as attraction by deception and three types were outlined. At first, the fragrances were thought of as a type of deception. Further research revealed that the Catasetinae, however, were not operating on deception. They attracted their pollinators by offering not food, but the chemicals found in their fragrances (Dodson 1962b). Allen and others had the idea that the insects gnawed on the lip and that this might provide some type of food, or by deception, make the bee think there was food. Stefan Vogel (1966) came to the conclusion that this theory was incorrect and along with Dodson and Frymire (1962a; 1962b) found that the pollinators were not gnawing the lip, but instead were brushing an area on the lip with specialized pads on the front pair of legs which extracted fragrance chemicals. The insect then flew out of the flower and

transferred the chemicals from the pads on the front legs to storage receptors on the back pair of legs. This behavior pattern appears to have led the Catasetinae to evolve a pattern of shooting their pollen very accurately onto their pollinator either when it enters or leaves the flower. This is a system which I understand is found in only one other genus among the Orchidaceae. The completely unrelated genus Listera R.Brown of tribe Neottieae employs a trigger system much like Catasetum; however, since it has perfect flowers, it also resembles the Mormodes and Clowesia system of not being receptive to pollination until after the pollinarium has been removed for a period of time (Ramsey 1950). This is a fascinating instance of parallel evolution since Listera is a northern hemisphere terrestrial which is certainly not pollinated by Euglossine bees.

In the Catasetinae each of the genera differs fundamentally in the device which triggers the pollinarium and the subsequent location on the insect of the pollinarium. The different systems are made possible by the location of the source of the fragrance which draws the insect into the correct position to receive the pollinarium. Dodson says that the presence of numerous and diverse pollinators has been of particular importance in the development of the many species found in the Orchidaceae. Adaptation to specific pollinators leads to the development of separate species from a common ancestor. Attracting specific pollinators establishes a group which does not interbreed with similar related species even though they may be in close proximity. This could be caused by development of different fragrances which attract different species of pollinators. Geographic or habitat separation may also lead to adjustment of a group of plants to pollinators different from those in their ancestral location.

While Dodson was working in Ecuador, Robert L. Dressler was beginning work in the field of pollinators in Mexico. He later moved on to Central America, especially Panama and Costa Rica. These two scientists and their collaborators did landmark work investigating and identifying the pollinators. Since the attraction of the pollinators was based on the fragrance of the orchids, logically the next step was to analyze the components of the fragrances of various orchids. This work was begun in the mid-1960's (Dodson & Hills 1966). Stefan Vogel (1966) identified the presence of specific scent organs in a number of orchids including the subtribes Catasetinae and Stanhopeinae. These organs, called osmophores, produce the chemicals which attract pollinators. Vogel found that the osmophores belonging to the Catasetinae and the Stanhopeinae differed from those of other orchids by a capacity to sweat out liquid drops of essential oil in specialized areas of the lips to attract the pollinators to the specific place which would insure correct placement of the pollinarium. The work is too extensive to examine in detail in the scope of this paper (see the Bibliography under Dodson, Dressler, Vogel and their collaborators). I do want to discuss parts which I think have special relevance as well as mentioning the work of others who have pursued research on the pollinators and fragrance analysis of Cycnoches. I shall discuss the work on the pollinators first. Dodson and Dressler identified the pollinators of the Catasetinae specifically as the males of species in three of the genera of the Euglossine bee tribe, the Euglossa, Eulaema and Euplusia.

In 1966, van der Pijl and Dodson published the first comprehensive work on the pollination of the Orchidaceae since Darwin (van der Pijl & Dodson 1966). Earlier, Dodson (1962a) had discussed the seven species in the genus Cycnoches as identified by Allen (1952). Based on the extensive work on pollinators done by Dodson, Dressler and others, van der Pijl and Dodson now began the move toward re-establishment as valid species of some of those reduced to synonymy by Allen. On pages 65 and 66 they said that Allen's treatment now appeared to be in error. Referencing work at that time still unpublished, it had been established that Cycnoches aureum (Allen's Cycnoches egertonianum var. aureum) was exclusively pollinated by Eulaema nigrita in Panama. Since the pollinator of Cycnoches egertonianum (type species) had been identified by Dressler as Euglossa ignita (van der Pijl & Dodson 1966 Page 184), they concluded that C. aureum was a separate species. They published photographs of the identified Eulaema nigrita on both the male and female forms of C. aureum and photographs of Euglossa ignita on both the male and female flowers of C. egertonianum (type). Based on then unpublished work by Robert A. Dressler, they found that four other forms of the Cycnoches egertonianum of Allen from Costa Rica and Panama were exclusively pollinated by specific species of Euglossa which do not visit other forms, even if they are in the same area. They concluded that this established these four varieties of Allen as unquestionably distinct species. These species are not identified in the text, or in the appendix on page 184, but I speculate from information in later sources, that these are probably Cycnoches guttulatum, C. pachydactylon, C. stenodactylon and either C. dianae or possibly C. densiflorum (Dressler 1993a).

In this appendix (van der Pijl & Dodson, 1966, p. 184), pollinators for several other Cycnoches species are identified, among which is Euglossa hemichlora as the pollinator of Cycnoches peruvianum, another one of the species which had been identified by Allen (1952) as synonymous with Cycnoches egertonianum. This is the sixth species to be re-established based on the identification of different pollinators for the species. The other pollinators listed are Eulaema cingulata for Cycnoches lehmannii; Euplusia superba for C. pentadactylon and Eulaema cingulata for C. ventricosum and for what Dodson continues to term C. ventricosum var. warscewiczii.

Dodson & Gillespie (1967, p. 12), while examining the size of the orchid family, pointed out that Allen in his revision (1952) was not aware that there were many variants in the Heteranthae which were only slightly different morphologically, yet they were valid species because they attract different species of pollinators with little or no hybridization among them. Dodson said that with further field work, it "seems reasonable to expect at least 50 species in the genus Cycnoches". Later, in the same source, page 28, in discussing causes of speciation through genetic shifts in parts of a widely distributed base species, Dodson said this appeared to be the case with Cycnoches ventricosum. He described the population from southern Mexico to mid-Nicaragua as "relatively pure", but a different form, C. warscewiczii, appears in southern Nicaragua, extending through Costa Rica to western Panama. He found it to be pollinated by the same species of bee that pollinates C. ventricosum, and that C. warscewiczii is "only slightly different [from C. ventricosum] but is horticulturally a much more pleasing plant with larger, fuller flowers". He continued that in eastern Panama, Colombia and Venezuela "a form called chlorochilon is found". He described it as even larger, but pollinated by a different bee. Then, in Ecuador, another form appeared, called Cycnoches lehmannii, which Dodson found to be "unfortunately nearly identical" to C. ventricosum. It is pollinated by the same species of bee which pollinates both Cycnoches ventricosum and C. warscewiczii. His conclusion is that Cycnoches lehmannii was considered a separate species because of the great geographical distance between northern Central America and Ecuador. He stated that it made better sense to consider the whole group as Cycnoches ventricosum distinguishing the others as subspecies or varieties. He does conclude that Cycnoches chlorochilon may be considered a separate species. The basis is not stated, but I assume it is because of the different pollinator involved. I find these observations extremely interesting, in that it seems the use of morphological differences as indicators of different species, now appear to be subordinated to the identification of the pollinators to establish species validity. I certainly agree with the place different pollinators hold in distinguishing species, particularly sympatric species which may be very similar but do not interbreed. Old fashioned geographic separation, however, still might have its place in speciation. I don't think Dodson was trying to eliminate it as a factor.

I have not been able to find any published work on fragrance analysis among the four species of the C. ventricosum complex. It is possible that despite the identification of the same pollinator for three of the four species, there may be differences in the chemical make-up of the fragrances. All four are very fragrant, but I think there is a detectable difference between C. chlorochilon and C. warscewiczii. When I had both species in bloom at the same time last season, I kept them side-by-side in my living room. I thought I could detect differences in their fragrances. I also observed that C. warscewiczii seemed to be fragrant for a longer period of time than C. chlorochilon, although the fragrance of the latter was stronger than the former. This observation could be a peculiarity of the two clones (or perhaps my nose), but it may also be a difference which might be worth more investigation with a large number of clones.

In the late 1960's, the fields of research which came into prominence were fragrance analysis and the study of the Euglossine bees. These two fields attempt to answer the question why Euglossine bees visit orchids which do not provide a food supply. As discussed above, the mechanism whereby the male bee extracts chemicals from the floral fragrances was well-known. The question was what use the bee made of these chemicals. This question is still not fully answered but there are a number of theories and progress seems to have been made in answering it (Dodson et al, 1969, Dodson 1970, Dressler 1967, Ackerman 1983, Lumer 1978, Williams 1982 and the numerous references in their bibliographies). I do not plan to go further into the discussion of this area since I believe I have covered the points of importance to this paper.

Fragrance analysis, because of its role in the question of speciation within Cycnoches, does bear some discussion. Research began in this area in the late 1960's. Gas chromatography analysis (Dodson & Hills 1966) revealed individual components which made up the fragrances of orchids. Scent organs in orchids, especially in the Catasetinae, were identified (Vogel 1966). Both of these areas have already been discussed above. Research on a number of genera has been done, but, for this paper, I am chiefly interested in work which involves Cycnoches and so shall not cover other genera. For those interested in more details than I provide, see Dodson 1970, Dodson et. al. 1969, Hills et. al. 1968, Williams 1982, Chong et. al. 1982, and the references listed in their bibliographies in addition to works already cited. There is one additional reference which I want to discuss in some detail. Dr. Katherine B. Gregg published a paper in 1983, which sought to examine the role of fragrance in speciation in Cycnoches. I wish to summarize her research and conclusions, because I think they are important to the question of the validity of species and their development. The research involved four different species of the Cycnoches egertonianum complex which were imported from Panama. The species involved were Cycnoches densiflorum, C. stenodactylon, C. dianae and C. aureum. I shall not go into detail about the methods and techniques employed that led to her conclusions. I would like to note that she found orchid fragrances, as a group, are varying mixtures of some 60 different chemical compounds. Individual fragrances are composed of from two to as many as eighteen, possibly more, of these compounds. The particular mixture gives the species its individual, characteristic scent. Some of these differences cannot be readily identified by humans, but they obviously are identifiable to the insect which is attracted.

She found that Cycnoches aureum had no identifiable fragrance and no odor compounds appeared in the gas chromatography analysis of any of the clones examined. This makes C. aureum unusual among the Cycnoches species. She offered two explanations, but the one she favored is that the flowers produce fragrance compounds which are not currently detectable. C. aureum has an identified pollinator, Eulaema nigrita (van der Pijl & Dodson 1966; Dressler 1968b), so there must be some chemical attractant for the bee. The other three species involved in the study, had very distinct and identifiable odors. Cycnoches dianae is described as having a sweet, lemony odor although two distinct chemotypes were identified among the clones examined with at least eight to ten different components present. Cycnoches densiflorum is described as possessed of a sharp pine scent although the components vary widely in their percentages among different clones. Some ten or eleven components appeared, not all of which could be chemically identified. Cycnoches stenodactylon is described as smelling sweet, reminiscent of rotten oranges. The fragrance components were distinctive among the three fragrant species. Dr. Gregg noted that the same distinctive compounds were found in both the male and female flowers of each of the species, insuring that the pollinator visited the correct female flower with pollen from the appropriate male flower. She also described each of the species, noting that they were easily distinguishable in morphological and color characteristics. Her methods of gas sampling verified that the lip area was the origin of the fragrance in all species examined. Dr. Gregg concluded that the noted differences among the four species justified their establishment as separate and distinct species.

To me, one of the most interesting results of her work was the identification of two distinct chemotypes in Cycnoches dianae. One type, representing about 35% of the clones she examined, lacked one of the chemical compounds which is normally a powerful attractant to some species of Euglossine bees. It has been well-established (see references at the beginning of this section) that presence or absence of compounds, can change the species of bee attracted. She speculated this chemical change could be a genetic mutation and that if the group which lacked the chemical should begin to attract a different pollinator from the normal group, it could be a beginning of the establishment of a separate species. She could find no morphological distinction to separate the groups based on color or lip form. She also identified different chemotypes among the clones of Cycnoches densiflorum examined. She reached similar conclusions in this case, speculating that, given the presence of appropriate bee species which might show a preference for a particular chemotype, the change could play an important role in further speciation. She concluded that the crucial relationship between the pollinators and floral odors made fragrance analysis a particularly useful tool in taxonomy.

Dr. Gregg's earlier work involving Cycnoches dealt with examination of ecological influences on the production of female flowers in the Catasetinae (Gregg 1973, 1975, 1976 and 1982). Dodson had first suggested that the production of female flowers, as opposed to male flowers in the Catasetinae, had an ecological basis (Dodson 1962a). He based this on field work in Ecuador with two species of Catasetum and one each of Cycnoches and Mormodes. He had observed that plants growing in full sunlight with adequate moisture were larger and generally produced female flowers. Less robust plants, growing in shade or lacking moisture, produced male flowers. He concluded that sunlight was the most important factor in the production of female flowers. To test this theory, he moved fifteen plants of Catasetum macroglossum which had bloomed with female flowers into a shadier location. These plants proceeded to produce male flowers. Of fifteen plants kept in full sun, thirteen continued to produce female flowers and two produced mixed inflorescences with both male and female flowers. Several large plants which had produced male flowers under shady conditions, were moved to full sun and they then produced female flowers.

To build on Dodson's earlier findings, Dr. Gregg began her research with importation in 1968 and 1969 of a large number of plants of several species of Catasetum and Cycnoches. I shall not go into the details of her research and control methods, but they appear to have been meticulous. The experiments in the initial research continued through 1973. She conclusively established that high light stimulated female flowering, but also theorized that other factors might play important roles (Gregg 1973 and 1975). In Gregg 1976, she published the continuation of her earlier work. Her further research found that light intensity interacted with plant size, health and nutrition to influence female flower production. This research used Cycnoches warscewiczii and Cycnoches densiflorum as its subjects.

Before moving on to final examination of the species, I wish to include some discussion of where Cycnoches fits into its subtribe, the Catasetinae. As has been previously outlined, the relationship of Cycnoches to Catasetum and Mormodes was recognized at an early stage. The three genera had been combined with closely related species of Cyrtopodium and Gongora to form the Catasetidae. Schlechter (1915), established the subtribe Catasetinae by separating these other groups into their own subtribe. Since then, the only major adjustment in the subtribe Catasetinae was Dodson's re-establishment of the genus Clowesia Lindley and erection of the genus Dressleria Dodson in order to clarify the genus Catasetum by removing the species with perfect flowers to separate genera (Dodson 1975). I refer those who are interested to Chase & Pippin 1990, Romero 1990 and Dressler 1993b, for details on the phylogenetic relationships among the Catasetinae. Here I only want to outline the cladogram which Romero arrived at in his analysis of the subtribe (Romero 1990, p 177; also found in Dressler 1993b, p.169). In the diagram (reading down), Romero places Cycnoches and Mormodes together as sister groups. Dressleria is placed as the next closest relative, with the three of them grouped as sisters to Clowesia. This entire group is then identified as sister to the two subgenera or sections of Catasetum. Dressler identifies the Cyrtopodiinae and Acriopsidinae as probably the closest related subtribes to the Catasetinae (Dressler 1993b).

In 1989, after a hiatus of almost fifty years, new species in the genus Cycnoches began to be published and described. In that year, Dr. C. H. Dodson, published four new species, Cycnoches bennettii, C. jarae, C. suarezii and C. thurstonorum. All four species are referable to the section Heteranthae with the first two identified from species collected in the Amazonian sections of Peru, and the last two identified from the eastern Amazonian portion of Ecuador. These species have forms and features which clearly distinguish them from Cycnoches peruvianum found in the same areas. Some of them have characteristics which are unique to the Cycnoches. Cycnoches bennettii has extraordinary color. The sepals and petals are ocher-red tinted greenish; the lip bluish gray-green with a pale rose mid-lobe. The column is a bright lemon-green and the ovary is purple-brown. These are colors and combinations not found in other Cycnoches.

Cycnoches jarae bears several branched inflorescences and a unique lip. The lip is united into a deep, cornucopia shape which resembles the shape of the lip of Galeandra more than that of any other Cycnoches species. The botanical drawings which accompany the type description seem to indicate that the flower may be resupinate, not non-resupinate as are those of all the other Cycnoches. If this is correct, a new mystery to the pollination story might be added if the flower is resupinate because the lip's orientation would affect the position of the pollinator . Since I was not able to locate photographs of these two species, I have depended on the type descriptions and drawings for these observations and interpretations. Cycnoches suarezii and C. thurstonorum resemble the more usual species of the Heteranthae, being differentiated by the shape and number of projections on the lip. One interesting additional difference of Cycnoches thurstonorum is that the color of the lip is described as orange.

In 1989, Die Orchidee published pictures of another new species of Cycnoches, but the species was not formally described and published until 1991 under the name Cycnoches herrenhusanum by Drs. Rudolf Jenny and Gustavo Romero. It is also referable to the section Heteranthae, but the sepals and petals are rounder and flatter than normal for species grouped in this section. The sepals and petals are an almost uniformly golden yellow color, another unique characteristic among the Heteranthae. The species was identified from specimens collected in the southern-most province of Colombia. According to Dr. Jenny (personal communication), the new species has subsequently been found in northern Ecuador. It is pictured in plate XIV-2.

In 1992, D. E. Bennett described and published another new species in the section Heteranthae, Cycnoches quatuorcristis. One distinctive feature of this species and the characteristic which gives it its name, is the four low ridges found on the yellow lip of the female flowers. The disc of the lip of the male flowers is markedly cupped, another distinctive feature of this species. The most recently described species of Cycnoches was found in the Brazilian state of Para and published by M.S. Campacci and P. Castro in 1993. It has been named Cycnoches manoelae. The authors describe the species as similar to C. thurstonorum Dodson, but distinguished from it by several differences in the sepals, petals and the lip. Cycnoches manoelae is also described as some 30 percent larger than Cycnoches thurstonorum. With the addition of these seven new species published recently, the total number of species in Cycnoches is brought to twenty-five at this time, six in the section Eucycnoches and nineteen in Heteranthae.

The remaining two parts of this paper deal with the hybrids involving Cycnoches and the American Orchid Society awards which have been granted to the genus and its hybrids. Hybridization and the awards are horticultural aspects, so before we move into these sections, I think we have to explore how the genus has been treated in horticulture. The treatment is not exactly the same as in botany and taxonomy. At the risk of oversimplification, I will say horticulture represents the "art" of growing plants as distinguished from the sciences of botany and taxonomy. I shall not discuss the culture of Cycnoches and the related genera. That is a subject entirely separate from the goals of this paper, and much has been written on it. The horticultural aspect which I want to discuss is the nomenclature that has been and is being used in horticulture to identify species. The historical confusion that dogged the genus Cycnoches has been the origin of the confusion of species and their names that plagues the hybrids and the awards. I chose the following books as sources: Veitch 1887-94, Williams, B.S. and H 1894, Sander 1927, Rentoul 1989 as well as the "standard" modern encyclopedias of orchid species, Hawkes 1965, Bechtel et. al 1986, Sheehan & Sheehan 1979 and 1994, and Pridgeon 1992. The orchid periodicals perhaps are even more representative. I chose to use the following four: the American Orchid Society Bulletin 1932-current, the Orchid Review 1893-current, the Orchid Weekly (originally named the South Florida Orchid Weekly) 1958-1967, and the Orchid Digest (1951-current). These sources cover the last one hundred years, the period which saw the increase and spread of orchid cultivation.

This discussion will be limited to the treatment of the large-flowered "Swan" or "Golden Swan" orchids as they have been called , the most widely-grown species in horticulture . At the beginning of the period under discussion (approximately the last one hundred years), Cycnoches chlorochilon was correctly identified in the books by Veitch and Williams ( op. cit.) . The latter provides an illustration of Cycnoches chlorochilon from Gardening World (although shown upside down). C. ventricosum is described by Williams ( op. cit.) but with the notation that it "sports" to C. egertonianum. This is probably a notation from the earlier editions of the book, which had first been published in 1865. Veitch ( op. cit.) does not describe Cycnoches ventricosum in his list of species, but he does refer to it in the description of C. egertonianum, relating the confusion which led to the conclusion that these two species were sports of each other. He does correctly state that the species are not the same and discusses the confusion of the female flowers of Cycnoches egertonianum with the male flowers of Cycnoches ventricosum. Neither of these books mention Cycnoches warscewiczii.

Sanders 1927 lists Cycnoches chlorochilon but the description is so general that it could be applied to any species of the C. ventricosum complex . The country of origin is given as Colombia which does not clarify the matter since C. chlorochilon was originally collected in Venezuela, although it is found in Colombia. For many years, these books were the main sources of information about species to the orchid-growing public as well as the commercial growers. This may be one of the reasons that Cycnoches chlorochilon became the name in general use for this type of Cycnoches. Another important fact seems to be that the true Cycnoches chlorochilon apparently fell out of cultivation somewhere around the early part of this century. This timing coincided with the entry of many United States citizens into Panama after independence was declared in 1903. Among these Americans were a number who were interested in orchids. American scientific institutions were also attracted to the area. One of the most attractive orchids found in Panama was the large-flowered "Swan Orchid" from the western, more accessible part of the country. The eastern section, near the Colombian border, is still almost inaccessible to this day. Therefore, the "Swans" being collected and exported were what we now know to be Cycnoches warscewiczii. As was discussed on pages 8 and 9 above, the identity of the Cycnoches warscewiczii of Reichenbach was not established until the early 1940's, so it is no small wonder that the Swan orchids were being identified as Cycnoches chlorochilon. They obviously were not Cycnoches ventricosum, and C. chlorochilon seemed to be the only other choice. In this period, the Cycnoches warscewiczii that appeared in publications and horticulture was based on the plant awarded in 1879 as mentioned above on page 9, and which had been pictured in Gardeners' Chronicle 12: 493 1879; Garden vol. 37: 225 1880, vol. 43: 217, 1893 and vol. 51: 172, 1897; and the Orchid Review (Unidentified Author 1917,McNab 1933). But this plant is really one of the Heteranthae which was flowering at the time of the award with both male and female flowers.

There are numerous examples of articles from the 1930's onward which discuss Cycnoches chlorochilon, but when there are pictures, the pictures are of Cycnoches warscewiczii Reichenbach f. (Unidentified Author 1932, Cope 1942, Dunn 1948, Wells 1950, Hawkes 1951, Scully 1951, Merkel 1951, Moore 1960, Sheehan & Sheehan 1974, 1979 and 1994, Rittershausen 1975, Cavestro 1977, Northen 1983, Batchelor 1983). It also seems that plants of Cycnoches warscewiczii were sold under the name Cycnoches chlorochilon. For example, Alberts & Merkel has an advertisement in the 1950 A.O.S. Bulletin , vol. 19(11) page 631, and the 1951 Orchid Digest, vol. 15(5) page 227, for Cycnoches chlorochilon. The picture is of Cycnoches warscewiczii. Bechtel et. al. 1986 do not even mention Cycnoches chlorochilon or C. warscewiczii. Cycnoches ventricosum is listed and described. The photograph, titled C. ventricosum, is of C. warscewiczii. Since this volume is supposed to be a basic reference for species, I think this is a great disservice to its readership. There were articles pointing out this error (Hawkes 1953, Hawkes 1959, Hawkes 1961, Butcher 1964, Northen 1973), but the misuse of Cycnoches chlorochilon, or even Cycnoches ventricosum var. chlorochilon continued. I was amused to read in Rentoul's 1989 revision, in the discussion of Cycnoches ventricosum, that he says the species has an "accepted variety chlorochilon [that] has sometimes been accorded the rank of separate species without any real right to the privilege...But the variety warscewiczii sometimes is passed off as it". There are more examples but I think the point is made with one last reference. Segars 1960 has a discussion of cultural requirements for growing Cycnoches chlorochilon among other species. He notes that several distinct kinds of Cycnoches have been introduced into horticulture under the name C. chlorochilon and makes no attempt to identify the species being grown. This view has led to mistakes in identifying plants both in hybridization and in the awards. The fact that the Registration Authority has considered Cycnoches chlorochilon an acceptable synonym of Cycnoches ventricosum and has more or less ignored Cycnoches warscewiczii all together further contributed to the confusion.

To date (September 1995), only thirty registered hybrids involving Cycnoches have been published . Ten of them are intrageneric Cycnoches hybrids; seven are Cycnodes ( Cycnoches X Mormodes); eleven are called Catanoches in registration, ( Cycnoches X Catasetum) but four of these used Clowesia as the " Catasetum" parent; and two are Monnierara ( Cycnoches X Mormodes X Catasetum). Appendix B lists the crosses, their registered parents, year of registration and the name of the registrant.

Hybridization within the Catasetinae, in general, is a rather recent field. In the 1950's, W. W. G. Moir, a man of protean interests in the Orchidaceae, was one of the first, if not the first, grower to become interested in hybridizing with the Catasetinae. In 1959 he registered the first Catasetum hybrid, Catasetum Grace Dunn, a cross of Catasetum warscewiczii and Catasetum roseum. Both of these species were transferred to the genus Clowesia by Dodson (1975), but the Registration Authority continues to register hybrids with Clowesia as Catasetum. In 1960, Moir registered a second hybrid, Catasetum Green Mansions ( Ctsm. viridiflavum x Ctsm. laminatum), the first true Catasetum hybrid.

Moir's third registered hybrid in the Catasetinae was the first of the Cycnoches hybrids, Cycnoches Hawaiian Lei. In Sanders', the parents are listed as Cycnoches egertonianum and C. dianae. Jenny et. al 1993 give the parents as Cycnoches aureum and C. dianae. The Registration Authority considers C. aureum as synonymous with C. egertonianum and may have changed the pod parent name. I could find no conclusive evidence on this point or photographs of the cross. From the parents, I would speculate that the appearance of the male flowers would be rather typical of those of the Heteranthae, with either a yellow or pink background color, possibly with small maroon spots. Although other intrasectional Heteranthae hybrids have been made, none have so far been registered. Of the remaining nine registered intrageneric Cycnoches hybrids, three are intrasectional Eucycnoches crosses, four are intersectional crosses between Heteranthae and Eucycnoches species, and the last two are second generation crosses of a Eucycnoches species with an intersectional hybrid parent.

At this point, I want to remind the reader that, unless specifically stated otherwise, the discussion of hybrids compares and describes the male flowers, because the female flowers of both sections are so similar. The male flowers of the two sections have different characteristics which make them attractive parents. The Eucycnoches species have the largest flowers and, generally, better substance than the Heteranthae species. On the other hand, the Heteranthae species are usually much more floriferous and many of them have brighter colors arranged in more interesting patterns.

The first group I want to discuss are the three intrasectional Eucycnoches hybrids. The first two are Cycnoches Chloroge ( C. chlorochilon x C. loddigesii) and Cycnoches Cygnet ( C. chlorochilon x C. haagii). Both were registered in 1970. While registered as crosses of Cycnoches chlorochilon, both of these almost certainly were made with Cycnoches warscewiczii and not the true Cycnoches chlorochilon. Among the differences between C. chlorochilon and C. warscewiczii are the shape of the petals and the manner that they are held in relation to the rest of the flower. Cycnoches chlorochilon has longer and relatively narrower petals than C. warscewiczii. Cycnoches warscewiczii holds its petals almost horizontally. Cycnoches chlorochilon has rather upswept petals paralleling the lateral sepals and framing the lip. These differences give Cycnoches warscewiczii a flatter, rounder flower. Cycnoches Chloroge shows the red-brown color of Cycnoches loddigesii on its column with a variable amount of red-brown marking, depending on the clone, on the sepals and petals. From the Cycnoches warscewiczii parent, it inherits relatively wide, horizontal petals and better substance than Cycnoches loddigesii. The cross has inherited good qualities from both parents and is a generally pleasing flower. This seems to be the most widely-available of the intrageneric Cycnoches hybrids. I have seen it offered from several sources. Cycnoches Chloroge was granted a Judges' Commendation by the A.O.S. when it was first shown in October, 1970, and was given the clonal name 'Mamasco'. This clone, or a division of it, was awarded a Highly Commended Certificate of 78 points in 1983. I congratulate the growers on maintaining the clone in cultivation for so long. As most growers of Cycnoches know, it is difficult to maintain a specific clone in growth for such a relatively long period.

The second hybrid registered in 1970, Cycnoches Cygnet, is intermediate in size between its parents. Cycnoches haagii is the smallest of the Eucycnoches species, with a natural spread of around 6.0 cms. compared to the other species which range from 9.0 to 14.5 cm in natural spread. There is a photograph of the cross published on page 598 of the A.O.S. Bulletin, Vol. 46(7) and in Batchelor 1983, page 834. The flowers show the flat shape of those of both parents with olive green to chartreuse color, displaying various amounts of tiny, red-black spots which come from the Cycnoches haagii parent. The distinctive two small, raised crests of Cycnoches haagii appear on the lip. The lip also bears the scattered red dots of C. haagii. I think it would be interesting to remake both these crosses with a true Cycnoches chlorochilon to see what the results would look like.

The third intrasectional Eucycnoches crosses is Cycnoches Bo Day Shus ( C. haagii x C. loddigesii). It was registered in 1987. Features of both parents are evident in the cross. The sepals and petals have the rather long, narrow shape of Cycnoches loddigesii, but with a natural spread approaching 10.0 cm, intermediate between the parents. The flatter shape of Cycnoches haagii has reduced the usually cupped shape of Cycnoches loddigesii. The reduction in cupping contributed to the size of the natural spread . Cycnoches haagii heavily influences the lip and color of the flowers. The lip is shorter and wider than the typical Cycnoches loddigesii lip and is a creamy-white color with a pale rose blush. The sepals and petals are an olive green color in contrast to the red-brown markings of Cycnoches loddigesii. All segments, including the column show the small red-brown spots of Cycnoches haagii, although Cycnoches loddigesii also has spotting on the lip. The solid red-purple column color of Cycnoches loddigesii did not come through in the hybrid.

All three of these intrasectional hybrids have generally pleasing flowers with some of the best characteristics of their parents. They possess many of the features which hybridizers hope for when they make a cross. The flowers are large, with good substance and usually good color. I think they are successful crosses.

The remaining six intrageneric Cycnoches hybrids are the intersectional crosses. The first of this group was registered in 1962 and named Cycnoches Tawny. The pod parent is listed as Cycnoches egertonianum (Heteranthae) and the pollen parent as C. chlorochilon (Eucycnoches). In reporting the registration, Alex Hawkes (1963) again corrected the pollen parent to Cycnoches ventricosum var. warscewiczii. It is the pod parent, however, which is of interest. Although Rivermont Orchids is listed as the originator and registrant of the cross, William Segars (1963b), described how this cross was made in December 1960. I have not been able to determine what precisely happened, but Mr. Segars' description of the cross is the only information I was able to find about C. Tawny. Mr. Segars described the Cycnoches egertonianum parent of his plants as having flowers "half again as large" as other Cycnoches egertonianum and the color is described as a "warm brown with an overcast of maroon to magenta." The resulting cross was described as "almost exactly that of C. chlorochilon in miniature. The natural spread of the flower is 2 3/4 inches" (about 6.8 cm). Its color is the yellow-green of chlorochilon, but with an overlay of mahogany." The mahogany overlay must have come from the pod parent, which is an unusually dark color form. The female flowers of C. egertonianum are usually green, so it would appear that Mr. Segars was either discussing a male clone in his article, or that the plant was not C. egertonianum, sensu stricta, but one of the other species which Allen (1952) had reduced to synonymy. It is not possible to determine this point from the article. The sex of the flower is never mentioned by Mr. Segars. The clone of Cycnoches egertonianum was given a varietal name of 'Kerimeri' by the Segars, but it was either never shown for award, or if shown, was not awarded. I have not been able to locate photographs of the parent or the hybrid.

Cycnoches Tawny is one of only two intrageneric Cycnoches hybrids to be used in a second generation cross. In 1976, Cycnoches Tawny was backcrossed to its C. chlorochilon parent and registered as C. Tres Colores. Monnier 1990 (page 1223) says that a cross has been made with C. Tawny and Cycnoches pentadactylon, but this hybrid has not been registered to date. Monnier ( op. cit.) describes Cycnoches Tres Colores as more influenced by Cycnoches chlorochilon, as might be predicted from the fact that it is 75% C. chlorochilon (or probably more correctly C. warscewiczii). It has larger, heavier flowers with the Eucycnoches shape. Cycnoches Tres Colores was the first intrageneric Cycnoches hybrid to be granted an A.O.S. flower quality award. The clone 'Comanche' was granted a Highly Commended Certificate of 79 points in April, 1976, in California. The award described the plant as bearing six flowers with a natural spread of 10.0 cm. The background color was light brown, with a green flush at the base of the petals and sepals. The lip was described as white with a central green callus. This combination of colors is the origin of the name "Tres Colores" which means three colors in Spanish.

In 1975, the second intersectional hybrid, Cycnoches Rip-van-Winkle was registered. As a cross of Cycnoches maculatum and C. chlorochilon, (most likely Cycnoches warscewiczii), the cross probably would resemble C. Chloroge in many respects, although possibly smaller. Cycnoches maculatum has a dark coloration like C. loddigesii and is one of the larger, less reflexed flowers of the section Heteranthae. I was unable to obtain either photographs or a description of the cross, but it should have been a good flower. Cycnoches Tres Colores which was discussed with its parent C. Tawny, was the third intersectional cross registered. In 1978, Cycnoches Pistachio Moon was registered as a cross of C. egertonianum and C. haagii. Cycnoches Pistachio Moon is one of the best of the intrageneric Cycnoches crosses in my opinion. It takes the round rather full shape of its Cycnoches haagii parent and combines this with the floriferousness of its Cycnoches egertonianum parent. The sepals and petals are a pistachio-green color, overlaid with chartreuse. The lip follows its Cycnoches haagii parent including the two small "pincher"-shaped crests. There is also a light sprinkling of dark maroon spots on the lip and a few on the sepals and petals. I find it a very pleasing flower. There have been two A.O.S. awards to the cross and they will be discussed in the next section.

Cycnoches Pistachio Moon is the other Cycnoches intrageneric hybrid to be used as a parent in a second generation cross. In 1984, Cycnoches Nova Luna was registered as a cross of Cycnoches Pistachio Moon and Cycnoches loddigesii. This cross has a larger flower than its C. Pistachio Moon parent, as would be expected since it is 75% Eucycnoches. It is described as green with a dark overlay of the red-brown or magenta color from its Cycnoches loddigesii parent. The hybrid retains the floriferousness of its Cycnoches Pistachio Moon parent, possibly reinforced by the Cycnoches loddigesii parent which, while 5 flowers are average, has been seen to produce twelve to fifteen male flowers on a really well-grown plant.

The latest intrageneric Cycnoches hybrid was registered in 1991 by Gene Monnier as a cross of C. pentadactylon and C. loddigesii. It was named Cycnoches Pentalodd in the old tradition in hybridization of combining the names of the parent species. In Monnier 1990, part 3, the hybridizer described the cross as producing "long sprays of large, nearly solid reddish flowers...[which] are a good average of the two parents in size and shape, with a terrific intensification of flower color in many of the seedlings." He goes on to say that this type of breeding is very under-explored and that much can be gained by both intra- and intersectional breeding. This cross highlights the best features of these intersectional crosses. This type of hybrid combines the floriferousness of the Heteranthae species with the larger flowers with better substance of the Eucycnoches species. These qualities of the two sections join to meet several of the goals of hybridization in general -- higher flower count, better substance, larger flowers and interesting colors. The features found in the species and hybrids of Cycnoches lead us to the discussion of the intergeneric hybrids which have sought to bring together the best features of Cycnoches with those of its relatives.

The most successful intergeneric hybrids of Cycnoches to date have been those with its most closely related genus, Mormodes. These intergenerics are called Cycnodes and seven have been registered. The first Cycnodes, and also the first intergeneric hybrid in the Catasetinae, was registered in 1961, the same year as the first intrageneric hybrid of Cycnoches, C. Hawaiian Lei. Cycnodes L. Sherman Adams was registered as a cross of Cycnoches chlorochilon and Mormodes wendlandii. In reporting the registration (Hawkes 1962), Alex Hawkes called this a "remarkable and handsome bigeneric cross between Cycnoches ventricosum var. warscewiczii (not C. chlorochilon as indicated) and Mormodes colossus (not M. wendlandii, as indicated)." This would begin intergeneric hybridization with as confusing a start as the genus. The breeder of Cycnodes L. Sherman Adams was Charles Slocum who was employed by Jones & Scully at the time, and they, not Slocum, are listed in Sanders' as the registrants. Slocum 1961 gives us an interesting account of how the hybrid came about. He states that the cross "was made more in the spirit of a joke than a scientific experiment" in November 1954. Slocum says that when the pod ripened, some of the seed was flasked, but the remainder was scattered on the top of the media of some Dendrobium plants. He estimated that of the six plants that were alive at the time he was writing, more than half came from the Dendrobium pots because the flasked plants were never strong. The cross attracted a lot of attention as the first intergeneric cross in a subtribe which had been little used in hybridization until then. The hybrid was granted an Award of Distinction on December 13, 1961, in Miami. Slocum ( op. cit.) described the flower as "caught in a group of opposing forces. The petals tried to reflex, as in Mormodes, but were restrained by the Cycnoches influence. The column tried to invert, but could not quite complete the turn, so it is twisted. The smooth Mormodes lip has acquired the callus ridge so prominent in this particular Cycnoches." This hybrid displayed valuable qualities and some of the characteristics that make Cycnodes crosses popular--the flowers were "quite long-lasting either on the plant or when cut", and they displayed colors not common in Cycnoches. Slocum described the colors as ranging from "straw-beige tones to the tawny bronze." The colors Mormodes can bring to the other genera of the Catasetinae are among its most attractive features in hybridization. Monnier (1990, part 3) calls Mormodes the most colorful genus of the Catasetinae, with several species having flowers in the red range, a color not found in the other Catasetinae, and relatively large flowers up to ten or more centimeters.

The second Cycnodes was registered in 1966 as a cross of Cycnoches chlorochilon and Mormodes colossus. It was named Cycnodes Ginger Snap. As reported by Alex Hawkes (1962) Mormodes wendlandii is taken as a synonym of Mormodes colossus (also see Bechtel et al. 1986). If this is correct, Cycnodes L. Sherman Adams would be the correct name for Cycnodes Ginger Snap. Since I have not been able to locate a photograph of Cynd. L. Sherman Adams for comparison, I cannot confirm this. In any event, Cynd. Ginger Snap is an attractive flower. The cross was granted an Award of Merit of 86 points in December, 1965, with the clonal epithet of 'Eureka', and a Highly Commended Certificate of 77 points in January 1985 with the clonal epithet of 'Discovery'. There is a color photograph of an apparently unawarded clone of the cross on the cover of the July, 1977, cover of the A.O.S. Bulletin, in conjunction with the article by William Cavestro on the culture of the Catasetinae (Cavestro 1977). The cross follows the resupinate character of Mormodes, but the column resembles the arching untwisted shape of Cycnoches. The shape of the sepals and petals is similar to the Cycnoches warscewiczii parent, although somewhat more cupped . The color is a tawny bronze overlaid on green with fine lines of red-brown spots radiating out from the center in some clones. The lip is flat like its Mormodes parent, but bears a small, narrow crest below the base of the column. The flower's natural spread is somewhat intermediate between the parents, around 9 cm. The award descriptions of this cross will be discussed in the next section of this paper. Cycnodes Thomas Krumpe was registered in 1969 as a cross of Cycnoches chlorochilon and Mormodes buccinator. I was not able to locate either a picture or a description of this cross, but Monnier, who says he has not seen the cross either, speculates that it should be similar to Cynd. Ginger Snap (Monnier 1990, part 3).

The next Cycnodes to be registered is, without a doubt, the most successful of the Cycnodes hybrids to date. It probably is at least one of the best, if not the best, of the Catasetinae crosses. The point depends in part on one's taste. It is Cycnodes Wine Delight, registered as a cross of Cycnoches lehmannii and Mormodes sinuata, in 1980. This cross has been granted six A.O.S. awards, five Awards of Merit, ranging from 80 to 89 points and, until 1994, the only First Class Certificate (91 points) in the entire Catasetinae subtribe. Two of the Awards of Merit were granted to plants in our judging region owned by John and Gloria Naugle. The clones were appropriately named 'Glo' and 'John'. The awards will be discussed in the next section of this paper. The most striking feature of this cross is its rich red tones, overlaid on a green base. The reds range from ruby to garnet with symmetrical venation which is of an even darker color. The red overlay is so intense that the base color is barely evident from the front. The form of the flower is rather flat with a varying amount of cupping, depending on the clone. The lip is generally flat ranging in color from pink to ruby red. The flower generally follows the resupinate nature of its Mormodes parent, with an somewhat arching column like its Cycnoches parent. With a natural spread ranging from 6 to 7.5 cm, it is not a very large flower, but it is striking. Some clones bear up to a dozen flowers per inflorescence.

Cycnodes Chocolate and Cherries was registered in 1987 as a cross of Cycnoches chlorochilon and Mormodes sinuata. This is an interesting cross to compare with Cycd. Wine Delight, since both have the same Mormodes parent. Cynd. Chocolate and Cherries has a more subdued color with an olive green base color, overlaid with red-brown. Gene Monnier, who made the cross, says the red was often muted to a brown or rosy mauve. He says he named the cross for the dark red lip against a striped brown background in some of the seedlings (Monnier 1990, part 3). The form of the flower is very similar to that of Cynd. Wine Delight. Gene Monnier ( ibid.) attributes the intensification of the red color of Cynd. Wine Delight to the almost solid dark green of the Cycnoches lehmannii parent. He says that the yellower coloration of Cycnoches chlorochilon brought out more of the color of the Mormodes parent and resulted in the muted color of Cynd. Chocolate and Cherries in contrast to Cynd. Wine Delight.

In 1990 and 1991, Gene Monnier registered the two most recent Cycnodes crosses, Cynd. Key Lime Pie ( Cycnoches haagii x Mormodes buccinator) and Cynd. Raspberry Yogurt ( Mormodes frymirei x Cycnoches pentadactylon). Cynd. Key Lime Pie is described by Monnier (ibid.) as "a lovely combination of chartreuse and creamy butter yellow." Key Lime Pie seems to be a perfect name for this combination. While I have not seen the cross, I wonder if the distinctive "pincher"-like crest from Cycnoches haagii was inherited by the cross. Many of the crosses of Cycnoches haagii, as mentioned above, did show this distinctive trait. Up to this point, all the registered Cycnodes had used one of the Eucycnoches species as the Cycnoches parent. Cycnodes Raspberry Yogurt took a new direction by using Cycnoches pentadactylon, one of the best of the Heteranthae species. I have not found a description or picture of the cross, but I would anticipate that the flower count should be good and from the name, I assume it would be a beautiful red-purple color perhaps overlaid on a cream background.

I think that all six, or seven if Cycnodes L. Sherman Adams and Cycnodes Ginger Snap are not the same cross, are successful hybrids in that they meet goals one is usually looking to accomplish in hybridization.

The second intergeneric group involving Cycnoches, both chronologically and arguably in importance, is known in registration as Catanoches, the combination of Catasetum and Cycnoches. Currently, for hybrid registration purposes, the Registration Authority does not accept Clowesia as a genus separate from Catasetum. For this reason, all the species used in hybridization which were removed to Clowesia (Dodson 1975) are still registered as Catasetum. For clarity's sake, I have chosen to use, in the discussion of the intergeneric hybrids, the name Clowenoches which was suggested by Rebecca Northen (1983) and used in Jenny et. al. 1993 for crosses involving Clowesia and Cycnoches. The correct name in registration Catanoches is reserved in this discussion for hybrids of the true Catasetum species and hybrids with Cycnoches. There are four Clowenoches to consider. Clowenoches Green Beret was registered by W.W.G. Moir in 1967 as a cross of Clowesia warscewiczii and Cycnoches ventricosum. The first flowering of the cross, before it was registered, was reported by Alex Hawkes (1967) as the cross of Catasetum warscewiczii and Cycnoches ventricosum var. warscewiczii. He reported that Moir described the flowers as having a lip minus any whiskers (so prominent in Clowesia), on a pointed jaw, with odd stripes and colors. He also reported "many flowers on an 8 to 10 inch spike," (20 - 25 cm) "having trouble deciding whether to go up or down." I was not able to find a photograph of this hybrid, but I speculate from the parents and the name that it would have been a nice shade of green.

Clowenoches Rebecca Northen was registered in 1973 as a cross of Clowesia rosea and Cycnoches chlorochilon. This is without a doubt the most popular of the Catanoches hybrids. It is still seen in cultivation from time to time. It is not to be confused with Catasetum Rebecca Northen which really should be Clowesia Rebecca Northen. Clowesia Rebecca Northen is a combination of the first hybrid registered in the subtribe Catasetinae, Clowesia Grace Dunn ( C. warscewiczii x C. rosea) backcrossed to its Clowesia rosea parent. I believe Clowenoches Rebecca Northen is a beautiful although small flower, with good features inherited from both parents. It is intermediate in size between its 2.5 cm Clowesia parent and its 11 to 13 cm Cycnoches warscewiczii parent. The fact that Cycnoches warscewiczii and not Cycnoches chlorochilon was the parent is confirmed in Rebecca Northen's article on "her" cross (Northen 1983). The parents are pictured in the article and it is clearly Cycnoches warscewiczii (although upside down) that is shown. Ms. Northen ( ibid.) describes the influence of Clowesia rosea as bringing the round shape with an "oddly mixed lip with pink tones and a pink fringe" to the flowers. She also states that it produced perfect flowers like Clowesia rather than dimorphic flowers like Cycnoches. She noted that the plant flowers from the middle or lower portion of the pseudobulb, an interesting combination of the parents since Clowesia has strongly pendant inflorescences from the bottom of the pseudobulb and Cycnoches bears arching inflorescences from the top of the pseudobulb. I note that the flower also seems to be resupinate like its Clowesia parent. The larger size and the rather prominent crest found on the lip are traits from its Cycnoches parent.

In 1980, Clowenoches Green God was registered as a cross of Cycnoches chlorochilon and Clowesia warscewiczii. As noted above, Moir's cross Clowenoches Green Beret was registered as a cross of Clowesia warscewiczii and Cycnoches ventricosum (probably C. warscewiczii). Clowenoches Green God most likely also used Cycnoches warscewiczii rather than the true Cycnoches chlorochilon. If this is correct, then Clowenoches Green God really is just the reverse cross of Clowenoches Green Beret. I was not able to find a photograph of Clowenoches Green God, but Gene Monnier says that this cross "closely mirrors" Clowenoches Green Beret (Monnier 1990, part 2). He also concludes that they may be the same cross, since " Cycnoches chlorochilon was considered a synonym of Cycnoches ventricosum."

Clowenoches Lunar Cheese is the last of the registered Clowesia x Cycnoches hybrids. It was registered in 1982 by Jones & Scully as a cross of Cycnoches haagii and Clowesia russelliana. A picture of the cross appears on page 1124 of the A.O.S. Bulletin as a part of Monnier 1990, part 2. It shows an interesting mixture of its parents. The sepals and petals are a nice chartreuse color with darker green lines radiating symmetrically from the center of the flowers, as appropriate to the name referring to the "green cheese" make-up of the Moon. Unlike its Cycnoches parent, the lip is long and narrow with a cupped shape, although it is sprinkled with the red-brown spots of Cycnoches haagii in the proximal part of the lip. As far as I was able to see, the distinctive "pinchers" crest of Cycnoches haagii is missing. Cycnoches haagii influence has opened and flattened the flower. While Clowenoches Lunar Cheese is not completely flat, its cupping is about intermediate between the flat Cycnoches haagii and the very cupped Clowesia russelliana. It appears from the column, that it is a perfect flower and shares the resupinate stance of the Clowesia. Monnier ( ibid.) notes another interesting feature from its Cycnoches parent. The flowers are produced from the top of the pseudobulb.

I think all three -- or four, if Clowenoches Green God is really different from Clowenoches Green Beret -- are successful hybrids. They meet several of the goals one is looking for when making a cross -- unusual or brighter colors, size, floriferousness and long lasting qualities in the flowers. They are perhaps not as showy as the Cycnodes hybrids, but Mormodes and Clowesia bring different qualities to the crosses.

The true Catanoches hybrids have not been as successful as the intrageneric or the other two intergeneric hybrids discussed so far, although they do have some good qualities. Gene Monnier, who has done the most work recently in hybridization in the Catasetinae, has written several articles about what goals he has been trying to accomplish in his hybridization program (Monnier 1987, 1989 and 1990 in particular). Perhaps the main goal he has worked toward is flower longevity. This quality is particularly important in the commercial production of orchids. He writes (Monnier op. cit) that the Catanoches generally have been disappointing in respect to flower longevity, most of them lasting only 7 to 10 days. There has been some success, however, in producing flowers without the extremely sensitive trigger mechanism of the true Catasetum. Flowers of Cycnoches and Mormodes have the longest life span, with the true Catasetum species having the shortest life span in the subtribe. Clowesia falls between the groups. Dressleria seems to be a good candidate to use, but so far no hybrids with Dressleria have been registered, although some have been made. The flowers of the Dressleria species are at least as long-lasting as those of Clowesia, if not more so. Dressleria flowers are borne on upright inflorescences, in contrast to the sharply pendant inflorescences of Clowesia. This upright stance could be a useful trait to incorporate into intergeneric hybridizing. On the other hand, Dressleria are not as floriferous as Clowesia.

Seven true Catasetum and Cycnoches hybrids have been registered, although several others have been made (Bergold 1960, Monnier 1989 and 1990, part 2). Four of them listed Cycnoches chlorochilon as the Cycnoches parent, two of them used Cycnoches haagii as parent and the seventh used Cycnoches pentadactylon, the only registered cross with a species from the Heteranthae section. For reasons already noted, it is probable that Cycnoches warscewiczii was the true parent of the first four crosses. Five of the seven Catasetum parents are from the large-flowered Catasetum expansum/ pileatum complex. The other two species used depart from this pattern and employ Catasetum tenebrosum and Ctsm. saccatum. Because of the similarities among some of the crosses, I am going to discuss these hybrids in groups based on parentage.

In 1978, Catanoches Crazy Creature ( C. chlorochilon x Ctsm. Orchidglade) was the first of the true Catanoches to be registered. The closely related Catanoches Midnight Jem ( C. chlorochilon x Ctsm. Susan Fuchs) and Ctnchs. Yellow Bird ( C. chlorochilon x Ctsm. pileatum) were registered in 1991. Catasetum Orchidglade is a primary cross of Ctsm pileatum and Ctsm. expansum. Catasetum Susan Fuchs is a second generation backcross of Ctsm. expansum and Ctsm. Orchidglade. Because these crosses are so closely related, the form of the flowers are quite similar. The wide range of flower color in the Catasetum pileatum/expansum complex from pale yellow to very dark red, plus the wide variation in spotting on the flowers of the complex, make interesting variations in the crosses, despite similarities in flower form. One observation, made by Gene Monnier about his cross Ctnchs. Yellow Bird, is that the seedlings produced two different flower forms (Monnier 1990, part 3). One group looked like the Catasetum pileatum parent, but the others had waxy, bright yellow flowers intermediate in form between the two parents. It appears that it is the second group that gave the cross its name, Yellow Bird. I recently obtained a plant of the first type described by Monnier ( op. cit.). The shape of the lip is quite similar to that of Ctsm. pileatum. The sepals and petals are more typical of the Cycnoches parent in that they are broader than those of their Ctsm. pileatum parent and they are also better spaced. From their shape and form it would appear that C. warscewiczii was the parent and not C. chlorochilon. I think it is a rather attractive flower. The color of my clone is more of a pale chartreuse than yellow except for the yellow crest and column.

The next registered hybrids I wish to discuss involve Cycnoches haagii with Catasetum expansum/pileatum parents. Catanoches Lemon Meringue was registered in 1982 as a cross of Cycnoches haagii and Catasetum Orchidglade. Catanoches Green Dragon was registered in 1986 as a cross of Cycnoches haagii and Ctsm. expansum. The flowers are rather similar as might be anticipated. Cycnoches haagii reduced the size of the flower, but it spread the sepals and petals into a fan-shape and flattened the lip. The lip shows the concave crest of the Catasetum pileatum/expansum complex rather than the raised "pinchers" of Cycnoches haagii. Ctnchs. Lemon Meringue is more yellow than the olive-green tones of Ctnchs. Green Dragon. Gene Monnier finds these five crosses have the most attractively-shaped flowers of the Catanoches hybrids with a combination of Eucycnoches and Catasetum expansum/pileatum parents (Monnier 1990, part 3).

The remaining two Catanoches are very different from the five discussed above. Catanoches Fantasy, registered in 1978, the same year as Ctnchs. Crazy Creature, is registered as a cross of Ctsm. saccatum and C. chlorochilon. This is the only Catanoches to receive an A.O.S. award, a Highly Commended Certificate of 75 points. Gene Monnier calls the flower "very interesting if not 'attractive' like the Ctsm. pileatum types" (ibid.). The flower is described in the award as having green sepals and petals overlaid with brown, a combination of the parents' colors. The lip is curled and green spotted with brown. The column is green with brown spots on top and yellow underneath. The awarded clone had a 10.3 cm natural spread, increased over that of its Ctsm saccatum parent by the influence of the Cycnoches parent.

The seventh Catanoches is a cross of Cycnoches pentadactylon and Ctsm. tenebrosum, registered in 1982. This cross uses parents completely different from the other six and was named Ctnchs. Penthouse. I was not able to locate either a description or a picture of the cross, but from its parents I would speculate that it would have long pendant inflorescences with dark flowers of good size. Like Ctnchs. Fantasy, it may also be more interesting than attractive.

The final intergeneric hybrids to be discussed are the tri-generic combination of Cycnoches, Mormodes and Catasetum. There are two of them and they are called Monnierara, named in memory of Gene Monnier's mother (Monnier 1990, part 3). Both of these hybrids used Cycnodes Wine Delight as the pod parent. The first was registered in 1990 as Monnierara Ruby Matrix (x Ctsm. expansum), and the second in 1991 as Monnierara Mary Rose. Monnier said Monnierara Ruby Matrix showed varying degrees of dominance from its parent genera (Monnier 1990, part 3), but after two years of blooming the cross, all of the flowers had the same form. They appeared to be bisexual, he said, but he had not been able, at that time, to mature a pod on any of them. He speculated that with the mixture of genes, the cross might be sterile. He also said ( ibid.) that the colors were not up to his expectations, but that at least a fair amount of the vibrant red color of Cycnodes Wine Delight had come through. A picture of the cross appears in Monnier 1990, part 3, page 1224. In this same paper, he mentioned his cross with Catasetum Susan Fuchs, which was still awaiting its first flowering at the time. He said he used a dark clone of Ctsm. Susan Fuchs, which might result in a cross with more color, or could turn out solid green as some other crosses had done with a similar parent.

After our examination of the registered hybrids, the question about the future in hybridizing with Cycnoches comes to mind. Gene Monnier has published the most about this question (Monnier 1990, part 3 in particular). Alex Capriles (1991) also discusses future hybridization possibilities and his ideas and those of Gene Monnier are in general agreement. As was mentioned in passing above, there are several other intrageneric and intergeneric hybrids with Cycnoches which have been made and many of them have been bloomed, but they have not been registered (Monnier 1989). Monnier has suggested that the most successful intrageneric Cycnoches hybrids may lay in intersectional crosses. Cycnoches Pistachio Moon, its progeny C. Nova Luna, and C. Pentalodd seem to bear out this idea. The recently described species contain some interesting qualities which I think might create some beautiful and interesting crosses, especially if crossed to Cycnoches haagii, C. loddigesii or C. lehmannii. Cycnoches bennettii is described as "peduncle brownish purple, sepals and petals ocher-red tinted greenish, the lip bluish gray-green, midlobe tinted pale rose, the column bright lemon-green, the anther pale cream-yellow, the ovary purplish brown." To my knowledge, this array of colors is unique among the Catasetinae. It would be interesting to see these colors introduced into hybridization, if they do come through in the progeny. This species also seems to be a good candidate for crossing with some of the large-flowered or intensely colored Mormodes. Cycnoches jarae has several fascinating characteristics which are atypical of Cycnoches and the other members of the subtribe. It is described as bearing "3-5 lateral racemes, frequently 1 or 2 with a short basal lateral branch...sepals and peals yellowish green with scattered, small brownish or purplish red spots,...lip clawed, lateral lobes subtuberculate, united below the middle, forming a deep tube, flattened above, margins irregularly lobulate, more or less incised." The Spanish description is more detailed and says that the basal branch bears from 3-6 flowers which open 8-10 days before the rest of the 22-25 flowers of the inflorescence. Here we have a branching inflorescence, a trait found nowhere else in the Catasetinae that I know of, plus a lot of flowers perhaps as many as a hundred. The tubular lip is also unique. Its shape is similar to the lip of Galeandra rather than that of any of the Catasetinae, although it is not entire as is that of Galeandra. Cycnoches herrenhusanum is a beautiful golden-yellow color with very flat, round sepals and petals for a member of the Heteranthae. These traits could be useful in both intra- and intergeneric hybridization. In fact, any of these species might introduce an entirely different look to Cycnoches hybrids .

Gene Monnier also suggests that crossing Cycnoches with a Catamodes might produce a much waxier flower, and that crossing a Cycnodes with a Catamodes might give very deep colors and heavy substance. These crosses would be Monnierara. Catamodes Black Magic 'Jem', AD, AM/AOS, might be a good candidate since it has been described as the darkest colored hybrid ever seen by the Judging Team. Its color is described as "solid mahogany black." It is a cross of Mormodes sinuata and Catasetum Orchidglade. Catasetum pileatum var. imperialis with its dark solid red color, would seem another good candidate. Crosses of Clowesia and Clowenoches might also offer interesting hybridization with Cycnoches and Mormodes and their hybrids. One must always remember, however, as Gene Monnier said, in discussing his color expectations for Monnierara Ruby Matrix, "the colors were not up to my expectations for this cross (a rather typical result)." Murphy's law applies to orchid hybridization also, it seems.

As published through Volume 26, number 2, Summer, 1995, of the Awards Quarterly there have been a total of sixty-two awards granted to Cycnoches and its hybrids by the American Orchid Society. Half of them, 31 in all, have been granted as awards to species of the section Eucycnoches. Four of this group have received two awards each, so we are in fact dealing with twenty-seven different clones. Twenty of these clones belong to what may be referred to as the Cycnoches ventricosum complex -- C. ventricosum, C. chlorochilon, C. warscewiczii and C. lehmannii. These are the first awards we shall discuss. It is this group which is the most confused. Appendix C lists all of the A.O.S. awards, grouped in the order which I have chosen to discuss them. Eighteen awards, granted to fifteen of the clones of the C. ventricosum complex, are awarded as Cycnoches chlorochilon. Two awarded plants are listed as Cycnoches ventricosum var. chlorochilon; one as Cycnoches ventricosum; one plant with two separate awards is listed as Cycnoches warscewiczii var. warscewiczii and the last award to this group is listed as Cycnoches lehmannii.

I have found only two articles which deal specifically with the awards to the Catasetinae, Capriles 1991 and Monnier 1994. I read these papers before I started examining the awards to Cycnoches in detail. I was amazed on first reading Gene Monnier's paper (1994), by his statement about awards to Cycnoches chlorochilon. To quote Monnier, "all of the plants awarded in the U.S. as C. chlorochilon are almost certainly C. warscewiczii." On the other hand, Capriles rather accurately describes the size of Cycnoches chlorochilon while making no comment on the fact that most of the awarded plants are misidentified. Monnier attributes many of the misidentifications to the fact that a large number of the awards are to female flowers. As has been noted earlier, it is notoriously difficult to identify the species on the basis of female flowers since so little information is available to differentiate among them. As also noted, there are species whose female flowers resemble each other. While I agree with Monnier's idea, I attribute the misidentification in large part to the fact that in general horticulture and, apparently even in the view of the Registration Authority, it is the name Cycnoches chlorochilon which is used for the "Swans", no matter what is correct botanically. As the reader will have concluded by now, I do not agree with this idea. I find that it is not that difficult to differentiate among the male flowers of the four species of what I choose to call the Cycnoches ventricosum complex -- C. ventricosum, C. chlorochilon, C. warscewiczii and C. lehmannii.

Cycnoches chlorochilon is distinguished by an almost clawless, sessile lip which is at least twice as long as it is wide. The normally upswept stance of the petals, to parallel and frame the lateral sepals and the lip, is also distinctive among the four species. Cycnoches chlorochilon has a somewhat cupped shape and the largest flowers among these four species.

Cycnoches warscewiczii can be distinguished by the round, flat shape of the flowers. The lip of C. warscewiczii is almost round, with a much more convex shape than C. chlorochilon. Cycnoches warscewiczii has a short claw to the lip, in contrast to the sessile stance of C. chlorochilon. The shape and color of the callus are different from C. chlorochilum. There is a noticeable indentation below the callus of C. warscewiczii in contrast to the rather flat area below the callus of Cycnoches chlorochilon. The depression at the base of the callus on C. warscewiczii has a darker color -- almost blackish green -- than that of C. chlorochilon. The petals of Cycnoches warscewiczii are less than twice as long as they are wide. The sepals and petals of Cycnoches chlorochilon are well over twice, sometime as much as three-times, as long as they are wide.

In contrast to the somewhat cupped form of male Cycnoches chlorochilon and the flat shape of the petals and lateral sepals of Cycnoches warscewiczii, the male flowers of Cycnoches ventricosum and Cycnoches lehmannii both have sharply reflexed petals and sepals, giving them a kind of "wind-blown" look. Both of them also have very distinct claws to the lip. The claw on the lip of Cycnoches lehmannii is even longer than the claw of Cycnoches ventricosum. The lip of Cycnoches lehmannii is more deeply convex than that of all the other species of this group with an almost bubble-shape, although flattened on the dorsal side. Cycnoches lehmannii is also the greenest of the four. Its color is frequently almost forest-green. There are other differences, but I think this summarizes most of the distinctive features of the male flowers.

The female flowers are more difficult to distinguish, but they do share some of the differences of their male counterparts. The female flowers of Cycnoches chlorochilon share the larger size, somewhat upswept petal stance of the male flowers. The female flowers of Cycnoches warscewiczii share the flatter, rounder shape of their male counterparts. The female flowers of Cycnoches ventricosum may not be quite as reflexed as the male flowers, but they are definitely reflexed. The sepals and petals of the female flowers of Cycnoches lehmannii are sometimes almost as flat as the male flowers of Cycnoches warscewiczii but they are smaller and their color is much greener. The lip of the female Cycnoches lehmannii shares the almost bubble-shape of its male counterpart. I caution the reader that these observations about the female flowers of this group do not mean that there are not female flowers of other species, such as from the section Heteranthae, which might not match these remarks. Identifying the species on the basis of female flowers alone is still a risky business.

I have worked out a key for the section Eucycnoches which is found at the end of this paper starting on page 66, in an attempt to help my readers identify the species of the section. I hope to complete this key to include the species of section Heteranthae. As previously mentioned, partial keys for the genus are found in Schlechter (1916), Allen (1952), Dressler (1993a) and others. I refer my readers to these sources for comparison. Plates I through VI illustrate the male and female flowers of the six species of the section.

We now move to an examination of the A.O.S. awards to this group. It has been difficult in some cases to analyze the awards and to determine if the plants have been correctly identified. In some cases, the difficulty is due to lack of a picture and/or an adequate description of the awarded flower. In some cases, the award is to female flowers which, as I have said, can be difficult to identify. In any event, from the available evidence it appears to me that of the fifteen awarded Cycnoches chlorochilon, nine are definitely Cycnoches warscewiczii. Two of the remaining six plants -- the earliest A.O.S. award to the genus, a Botanical Certificate to the species in 1944, and the Certificate of Cultural Merit in 1980 to the clone 'Meri' -- lack sufficient information to identify the correct species. I feel that they are both probably Cycnoches warscewiczii. I base this on the fact that Cycnoches chlorochilon probably was not in cultivation at the time the Botanical Certificate was awarded. In the case of the clone 'Meri', it was owned by the Segars, who, as pointed out above, were known to have been growing Cycnoches warscewiczii not Cycnoches chlorochilon (Segars 1960, 1962 and following). Two of the awards, the clones 'Anita Bergold' and 'Green Jade', are to plants bearing all female flowers. After examining the award descriptions and photographs, I believe that the clone 'Anita Bergold' probably is C. chlorochilon and the clone 'Green Jade' is C. warscewiczii. Using the criteria described above, the photographs of the two clones clearly show the differences in the sepals, petals and lip which distinguish the male flowers of the two species.

This brings us to the last two of the fifteen " Cycnoches chlorochilon" awards, the clones 'Giant' and 'El Hatillo'. I have not been able to obtain a picture of either one of these clones, so I have had to rely on the award descriptions and measurements in them. I looked at the proportions of the measurements given for the sepals, petals and lip of Cycnoches chlorochilon 'Giant'. The length is in all three cases some three times the width, which makes the proportions of the segments typical of C. chlorochilon rather than C. warscewiczii. Based on this admittedly speculative, but I believe reliable, evidence, and despite the rather early date of the award, 1973, I think it is probable that the clone 'Giant' is Cycnoches chlorochilon. When the A.O.S. Awards Registrar in West Palm Beach is able to obtain the slide for me, I may be able to resolve this question.

The last of this group awarded as Cycnoches chlorochilon is the clone 'El Hatillo'. No award photograph was made of this plant. I examined the proportions of the segments as I did in the case of the clone 'Giant'. These measurements, in addition to the fact that the plant was awarded at a show in Venezuela, lead me to the conclusion that it is almost certainly a true Cycnoches chlorochilon. Its description, measurements and size, plus the fact that there are only two flowers, lead me to the conclusion that they are female flowers, although the sex of the flowers is not mentioned. I think this would have been an important piece of information to include. In general, I believe that the sex of the flowers should be determined before judging any flower from the subtribe Catasetinae. This information has an important bearing on how to judge any of the dimorphic members of the Catasetinae, not just the Cycnoches. This is not my original idea. Gene Monnier (1994) stresses the point that the sex of the flower must be determined or the plant may not be fairly judged.

As yet, we have only three probable A.O.S. awards to the true Cycnoches chlorochilon out of these fifteen plants awarded under this name. What about the two clones awarded as Cycnoches ventricosum var. chlorochilon, 'Cherimoya' and 'Peter Micha'? Unfortunately, they are neither Cycnoches ventricosum or Cycnoches chlorochilon. Both of them are clearly Cycnoches warscewiczii. Next, let us look at the awarded Cycnoches ventricosum 'Riopelle'. It is clearly another clone of Cycnoches warscewiczii. In my opinion, we now have fifteen Cycnoches warscewiczii all awarded as Cycnoches chlorochilon, and three probably true Cycnoches chlorochilon, two with female flowers and one with male flowers. This brings us to the two awards which have been granted under the name Cycnoches warscewiczii var. warscewiczii. This plant, with the clonal name 'Jan's Swan', was awarded a Certificate of Botanical Merit in 1975 on a blooming of all female flowers. The award description stated that this is the "true" C. warscewiczii var. warscewiczii and not the commonly awarded Cycnoches chlorochilon type. Suspicions about the validity of the species might come to mind based on the fact that there are purple-brown markings about the column, a feature not found on either the male or female flowers of any of the four species of the Cycnoches ventricosum complex. On page 9 above, I recounted the description by Rolfe (1909d) of the R.H.S. award in 1879, to a plant described as Cycnoches warscewiczii Rchb. f. The plant was blooming at the time with one inflorescence of female flowers and one of male flowers. The male flowers are definitely of the section Heteranthae and, as best can be determined from the plate, may be Cycnoches egertonianum, or one of its complex. They are certainly not what eventually was established to be Reichenbach's Cycnoches warscewiczii. But to requote Allen (1952), "error breeds error." This award from 1879 may be the origin of the idea that 'Jan's Swan' was Cycnoches warscewiczii. In the case of 'Jan's Swan', however, we have a resolution to the question of its correct species. Three years after the first award, Cycnoches warscewiczii var. warscewiczii 'Jan's Swan' was presented a second time for judging. This time the plant bore male flowers. It was granted a Highly Commended Certificate of 77 points, but from the award photograph, C. warscewiczii var. warscewiczii, 'Jan's Swan' may be identified from what can be seen of the lip and claw structure, as probably Cycnoches pentadactylon!

We have now covered nineteen of the twenty awarded clones of this group and we may conclude that technically Cycnoches warscewiczii may still be eligible for a Certificate of Botanical Recognition, since it has never been correctly awarded under this name. The true Cycnoches warscewiczii have all been awarded as Cycnoches chlorochilon and the one awarded plant described as Cycnoches warscewiczii is probably Cycnoches pentadactylon. But, let us return to our review of the awards by considering the last of this group of twenty plants of the Cycnoches ventricosum complex. In 1980, a Certificate of Botanical Recognition was granted to Cycnoches lehmannii 'Verde Cal'. After the discussion so far of distinguishing characteristics among the species of the Cycnoches ventricosum complex, a good look at the award slides shows us that at last we have an award to a true male Cycnoches chlorochilon. Unfortunately, it was certified as Cycnoches lehmannii and so the award stands today in the record. A strange ending to our examination of the awards to this group of plants. These awards are, if possible, more confused than the botanical history of Cycnoches.

I wish to say that, although the validity of the species names of these awards are, as I think I have proved, not correct in most cases, I do not think that the awards, except for the Certificate of Botanical Recognition, are undeserved. The plants which earned Certificates of Cultural Merit, the Awards of Merit, the Judges' Commendations, and the Highly Commended Certificates, were worthy of the awards. I do wish they had been made under the correct names. It would certainly make judging of future members of this group much easier.

Examination of the rest of the Awards is much more conventional. Those to three clones of Cycnoches haagii and four clones of Cycnoches loddigesii round out the awards to the section Eucycnoches. The plants are all correctly identified, unlike their sectional relatives discussed before. None of the three awards to Cycnoches haagii are flower quality awards. The first award to Cycnoches haagii was a Certificate of Botanical Merit of 81 points to C. haagii 'Deerfield' granted in July 1975. Cycnoches haagii 'Heritage' was granted a Certificate of Cultural Merit of 81 points in September of the same year. One year later, Cycnoches haagii 'Miami' was awarded a Judges' Commendation. No more awards have been made since 1976 to Cycnoches haagii. All three plants are beautifully grown and flowered examples of the species. The clone 'Miami' has no description or measurements, except that it is commended for "gorgeous, brilliant, rich-green color, outstanding, rich, velvety color and texture, contrasting brilliantly with white lip spotted red." All are male flowers, although this is not mentioned in any of the descriptions. Of the four awards to Cycnoches loddigesii, one is a Highly Commended Certificate of 76 points to the clone 'Marion', granted in 1984. The first award to Cycnoches loddigesii was a Certificate of Botanical Merit of 80 points, granted in 1968 to the clone 'Lorraine'. The other two awards to Cycnoches loddigesii are both Certificates of Cultural Merit. In 1968, the clone 'Caripe' was given 82 points, and the latest award was to the clone 'Emiria' granted in Venezuela in 1991 with a point score of 86. The last plant had a spectacular display of 50 flowers and 30 buds on eight inflorescences.

The section Heteranthae has received a total of fifteen awards to thirteen different clones. Cycnoches aureum 'Amarillo Cal' received a Certificate of Botanical Recognition in 1979. In 1987, Cycnoches egertonianum var. dianae 'Woodpoint' was granted a double award of a Certificate of Horticultural Merit, 83 points, and an Award of Merit, 83 points, in our own judging center in Tampa. A Certificate of Botanical Recognition was granted to Cycnoches maculatum 'Mackay' in 1979. All of these plants are beautiful examples of their species and bore male flowers.

Cycnoches egertonianum is the most awarded species of the section, with five awards -- a Certificate of Botanical Merit, one Judges' Commendation, one Certificate of Cultural Merit, and two Awards of Merit. These awards were given to an interesting variety of clones. The first award granted was C. egertonianum 'Lorraine' which received a Certificate of Botanical Merit of 85 points in 1961. It is described as bearing one inflorescence with four lime-green pistillate flowers and one small staminate flower spotted red. There was a second inflorescence on another pseudobulb with fifteen buds. No measurements are given. The female flowers may have been the feature which attracted the Judges' attention to the plant.

The second award was Cycnoches egertonianum 'Chadds Ford' which received a Certificate of Cultural Merit of 81 points in 1968. Again all of the fourteen flowers on four inflorescences are female flowers.

One month after the C.C.M. was awarded to the clone 'Chadds Ford', Cycnoches egertonianum 'Mickey' was granted a Judges' Commendation for "the rich magenta color of the sepals and petals, and for the increased width of the sepals and petals." No other information is given. The award slide is not very helpful. The slide shows only two of the flowers completely, one front view and one, on the opposite side, from the back. The flower is indeed a beautiful evenly-colored magenta with relatively wide, flat sepals and petals, not at all reflexed as is the norm for Cycnoches egertonianum. What little can be seen of the column appears to be rather long and arched like that of male flowers, however it is possible that these flowers are hermaphroditic of the form which is closer to the male flower than the female flower (see discussion on page 7). They are certainly not typical male or female flowers of Cycnoches egertonianum.

The fourth clone, Cycnoches egertonianum 'Cabrillo', was granted an Award of Merit, 81 points in 1979. This is an extremely interesting plant because it had 25 male plus one hermaphroditic flower on one inflorescence and a second inflorescence with two female flowers. This plant would have driven Lindley out of his mind had it appeared in his day (see discussion on pages 5 and 6). The award description of this plant is very detailed, which for a plant of the complexity of this one, is to be highly commended. I recommend it to anyone interested in an example of concise inclusion of information, although I would not want to try to fit it on the standard A.O.S. award form. The color of the male and the hermaphroditic flowers is described as almost black, although the perfect flower is said to have green striations on the lip and petal. Its size of 2.7 cm is intermediate between the male (1.5 cm) and female flowers (6.0 cm). I would note that the color is exceptional for Cycnoches egertonianum, sensu stricta. The type description in Bateman t. 40 of the species indicates dark color, but most of the Cycnoches egertonianum clones are not of such solid color.

The last of the five awarded clones is Cycnoches egertonianum 'Turrialba' AM/AOS, 86 points, which was awarded in 1989 at a show in San Jose, Costa Rica. The plant is described as having 44 flowers and 13 buds on two male inflorescences and three flowers on two female inflorescences. The male flowers are described as having a light green background with a heavily overlaid maroon color on the sepals and petals and a bright green lip. This is a very impressive display of flowers, although Cycnoches egertonianum frequently carries 25 or more flowers on an inflorescence. The size of the flowers is also impressive. The male flowers are said to have a natural spread of 2.7 cm and the female flowers with a natural spread of 8.2 cm. Comparing these sizes to Cycnoches egertonianum 'Cabrillo', the male flowers are some 40% larger and the female ones are more than 30% larger. The flower size made me wonder if perhaps this plant is incorrectly identified. The male flowers also seem to be closer together than normal for Cycnoches egertonianum. Cycnoches guttulatum, C. stenodactylon and C. pachydactylon are all found in Costa Rica. As discussed above, Allen 1952 places all of these species in synonymy with Cycnoches egertonianum. I tried to key out the flowers from the description and the unfortunately rather poor award slide using Dressler (1993a). I could not determine which species this plant is, but I do not feel certain that this is Cycnoches egertonianum, sensu stricta. Whatever the answer to this question, the plant richly deserved its Award of Merit.

The second most awarded species of the section Heteranthae is Cycnoches pentadactylon. It has received four awards (not counting the J.C. and H.C.C. granted as Cycnoches warscewiczii var. warscewiczii which were discussed above). The first award was a Certificate of Horticultural Merit of 82 points, granted to C. pentadactylon 'Spotted Swan' in 1985. Next were two Certificates of Cultural Merit granted to the clones 'Kilworth' (86 points) and 'Dr. John Doherty' (89 points). Cycnoches pentadactylon 'Dr. John Doherty' was simultaneously granted an Award of Merit of 87 points, the only flower quality award to Cycnoches pentadactylon and one of only two flower quality awards to strictly male flowerings in the section Heteranthae. As mentioned above, Cycnoches dianae 'Woodpoint', which was awarded in our own region, is the other clone to receive a flower quality award. Cycnoches pentadactylon is probably one of the most easily recognized species of the Heteranthae section. It also has the largest flowers. The clone 'Dr. John Doherty' had a natural spread of 10.0 cm and a vertical measure of 9.2 cm. While these dimensions are some 20% greater than typical, the average C. pentadactylon still is around 7.0 cm or more in natural spread.

I have saved the discussion of Cycnoches peruvianum var. album, the remaining award in the section Heteranthae, for last. Cycnoches peruvianum var. album 'J.E.M.' was granted a Certificate of Botanical Recognition in 1989, with eight flowers on two inflorescences. The award description states it is a rarely seen album form which is "remarkable for clear apple green color of sepals and petals and attractive arrangement of the inflorescence." Although the award description makes no mention of the fact, the flowers are clearly female flowers. Normal male flowers of Cycnoches peruvianum have purple or reddish-brown markings on the sepals and petals, with the typical long, many-flowered inflorescence of the section Heteranthae. I have not been able to obtain a slide of female flowers of Cycnoches peruvianum, but descriptions of the female flowers are found in Dodson & Dodson 1982 and Dodson & Bennett 1989. In both places, the female flower of Cycnoches peruvianum is described with grayish green sepals and petals, a white lip with a gray-green apex and a bright green column. This description fits this plant very well - see Plate XIII-2 for a picture of the plant. I won't dispute that an album form of Cycnoches peruvianum may exist, or that this plant might be such a form. I do feel that this blooming does not prove it. The flowers appear typical for a normal female Cycnoches peruvianum. I would wish to see a male blooming of this plant to determine if it is really an album form. Only a male blooming with clear green or white flowers would demonstrate that. I do not feel that a Certificate of Botanical Recognition is appropriate for this plant, although it is a fine flowering. First, as I have mentioned earlier, awards to female Catasetinae in general are difficult at best, unless they are flower-quality awards. While sex of the flower might be considered as immaterial to quality, I do think that the sex of the flower is the first point to be determined before any flower of the Catasetinae may be judged fairly. This completes the awards which have been granted to the species of Cycnoches, a total of 45 awards, sixteen of which are flower quality awards.

Of the ten intrageneric Cycnoches hybrids which have been registered, four have received A.O.S. awards. Cycnoches Chloroge received a Judges' Commendation in 1970 and a Highly Commended Certificate of 78 points in 1983. Both awards were to the clone 'Mamasco'. When the J.C. was published in the Awards Quarterly, Vol. 2, No. 3, July 1971, the cross was pictured on the cover, since it was a first of its kind and the hybridizer-grower was commended "on a refreshing and unusual new hybrid." The award included no measurements, but its form and color resemble the C. loddigesii parent, while the C. warscewiczii parent widened and brought the petals into a more horizontal plane. The flowers are clearly male, although no mention is made of this. Thirteen years later, the clone was shown again with two flowers. This time it received a Highly Commended Certificate of 78 points. The color is described as "deep apple-green with light mahogany flush on sepals and similar striation of sepals...heavy substance." It is remarked that "sparse number of flowers decreased the score." One look at the flower shows it to be a female, not a male flower, just from the form of the column (see plate VII-1). The larger size and heavier substance are also indicators of the sex of the flower. It is normal for female inflorescences to have fewer flowers, so the failure to recognize the sex of the flower may have decreased the score enough to deny the plant an Award of Merit.

The second intrageneric hybrid to be awarded was Cycnoches Tres Colores 'Comanche' which received a Highly Commended Certificate of 79 points in 1976. There is no award photograph and I was not able to obtain a photograph of the cross from any other source. It is a cross of one intersectional parent, C. Tawny with C. chlorochilon (probably C. warscewiczii). The cross has only six flowers on one inflorescence with a natural spread of 10.0 cm. The color is described as a background of light-brown with a green flush at the base of the petals and lateral sepals. The lip is said to be white with a central green callus. The double dose of Eucycnoches from the grandparent on the pod side and the parent on the pollen side certainly increased the flower size, but decreased the flower count, unless of course these were female flowers. No reference to the sex is given and no dimensions or description of the column is included, so this is impossible to determine without a picture.

The next Cycnoches intrageneric hybrid to be awarded was C. Pistachio Moon. In 1981, Cycnoches Pistachio Moon 'The Prophet' was granted a Highly Commended Certificate of 77 points. It bore four flowers and one bud on a single inflorescence. The sepals and petals are described as medium green with a bronze overlay and stripes appearing dark olive, with sepals darker than the petals. The lip was "polished-pink ivory" with waxy substance. The natural spread was 9.2 cm with column length of 1.5 cm. An examination of the award photograph (Plate VIII-2), along with the clues from the description, clearly show these are female flowers, a fact that is not mentioned. The flower count is in fact very good for a female inflorescence and if this was not noted, it may have, as in the case of Cycnoches Chloroge 'Mamasco', decreased the score the clone received. The potential of this intersectional cross of C. egertonianum and C. haagii was shown when Cycnoches Pistachio Moon 'Victor' was shown and awarded a Certificate of Cultural Merit of 82 points in 1984 (Plate VIII-1). This time the award states there are fifty male flowers and six buds on three inflorescences, all originating from the same pseudobulb. This demonstrates what can be produced by introducing the floriferousness of the Heteranthae to the larger size of the Eucycnoches. The natural spread is given as 7.0 cm which is of course considerably smaller than the previously awarded clone 'The Prophet', whose flowers were female.

The last intrageneric Cycnoches to be awarded is Cycnoches Bo Day Shus 'Jadeite Swan' which received a Highly Commended Certificate of 79 points in 1992 (Plate VII-2). This Eucycnoches cross of C. haagii and C. loddigesii combines the best features of both parents. It bears some resemblance to Cycnoches Chloroge, but is slightly smaller and has the distinctive crest of Cycnoches haagii on the lip . I personally find it to be a beautiful flower. The flowers appear to be male.

The most successful intergeneric hybrids of Cycnoches have been the Cycnodes. Four of the seven registered hybrids have received A.O.S. awards. Cycnodes L. Sherman Adams was granted an Award of Distinction in 1961 when it was first shown, in part because it was the first intergeneric hybrid in the subtribe Catasetinae. As I have stated in the section on the hybrids, I have not been able to locate a photograph of the cross and no information is provided in the award description. The appearance of the cross as described by the hybridizer was given in the section on hybrids. Cycnodes Ginger Snap, which may in fact be the same as Cycnodes L. Sherman Adams, as was also previously said, has earned two A.O.S. awards. The first clone to be awarded was Cycnodes Ginger Snap 'Eureka' which earned an Award of Merit of 86 points in 1965. I was not able to obtain a copy of the award slide, but the description sounds different from that of the clone which appeared on the cover of the issue of the A.O.S. Bulletin containing Cavestro 1977. The award description says there were six tawny brown flowers of very heavy substance with large, flat lips with yellow centers. The natural spread is given as 3 and 1/8 inches (approximately 7.7 cm). No dimensions were given for the clone on the cover of Cavestro 1977, but the appearance of the flower is different, being covered with lines of tiny dots radiating out from the center of the flower. In 1985, a second award was granted to Cycnodes Ginger Snap. The clone 'Discovery' was granted a Highly Commended Certificate of 77 points. This clone is described as also having six flowers with the sepals and petals a greenish brown base color, radiating to light reddish brown on the margins. The natural spread is 9.2 cm, considerably greater than that of 'Eureka'. The lip is light brown with darker reddish brown on the margin and base. The lips of both awarded clones are almost round, based on the dimensions given. These two clones are noticeably different from the clone pictured on the A.O.S. Bulletin cover of Cavestro 1977. Neither of the awards mention any lines of small dots which are evident on the unawarded clone. The lip of the clone on the A.O.S. Bulletin cover is also quite different in shape. It looks at least twice as long as it is wide. I speculate that the awarded clones are both female-influenced flowers, whereas the clone on the A.O.S. Bulletin cover may be more male-influenced. It has only recently been recognized that Mormodes does have flowers which may lean more to female expression or male expression as well as intermediate forms (Monnier 1992). Gene Monnier says there seem to be two different types of flowers produced in the crosses with Mormodes. One type produces numerous flowers which tend to have their lips aligned with the flower stem, similar to the Mormodes parent. He terms these flowers "male-dominant." The other type of flower tends to be larger, but there are fewer per inflorescence. The lips are uppermost in these flowers as those of the Cycnoches parent. Monnier terms these "female-dominant" flowers (Monnier 1990, part 3). Some form of sexual variation of this nature may account for the differences in the Cycnodes Ginger Snap clones. The flower of Cycnodes Ginger Snap pictured in Monnier 1993, part 3, is similar to the two awarded clones.

In award terms as in most other respects, Cycnodes Wine Delight is by far the most successful hybrid of Cycnoches to date . This cross has been granted five Awards of Merit ranging from 81 to 89 points and, until 1994, it held the only First Class Certificate granted to any species or hybrid in the Catasetinae. The awarded clones are listed in Appendix C and I shall not discuss them individually. For illustration, I do want to provide some of the statistical ranges of the awarded clones. The natural spread of the awards range from 6.1 cm ('Orchid World' AM 81 points) to 7.4 cm ('John' AM 81 points). The colors are variously described as shades of deep red to garnet and mahogany, with deep pink or red lip color. There are generally radiating, symmetrically arranged, darker colored lines covering the sepals and petals. Flower count ranges from a high of twenty-nine on two inflorescences ('Mendenhall' AM 89 points) to three flowers and two buds ('J.E.M.' FCC 91 points). The sepals and petals are relatively flat, although some cupping appears. The lips are generally flat, with the exception of 'Mendenhall' whose lip is somewhat convex. Even unawarded clones of this cross which I have seen, are worthwhile additions to any collection. The latest award to this cross was granted in our region in 1994 ('John' which belongs to the Naugles, as did the clone awarded with the epithet of 'Glo'). The FCC clone 'J.E.M.' is shown in Plate XV-2.

The last Cycnodes I shall discuss is Cycnodes Chocolate and Cherries which was granted an Award of Merit of 83 points in our judging region in 1991. It belongs to the Reeses of St. Petersburg and like Cycnodes Wine Delight is a Mormodes sinuata cross, with the other parent listed as Cycnoches chlorochilon. As mentioned in the section on hybrids, the hybridizer Gene Monnier felt that this cross did not come up to his expectations for color. It is, however, a beautiful flower, even if the color is not as intense as that of Cynd. Wine Delight. It shows the flat lip of Cynd. Wine Delight, and the natural spread of 6.1 cm is the same as for the awarded clone of Cynd. Wine Delight 'Orchid World'. I have also seen unawarded clones of this cross and they would be a nice addition to any collection.

The only other intergeneric Cycnoches hybrid which has been awarded is Catanoches Fantasy 'Scorpio' which was granted a Highly Commended Certificate of 75 points in 1978. According to the A.O.S., there is no award slide, but the description indicates a rather heavy influence from the Ctsm. saccatum parent. The plant bloomed from the bottom of the pseudobulb with eleven flowers on the inflorescence. The lip is described as curled. The color of the sepals and petals is described as green with an overlay of brown. The lip is green with brown spots on top and yellow underneath. The heavy substance of the flower probably did come from the Cycnoches parent along with the base green color. The natural spread is given as 10.3 cm which could be attributed to the influence of its Cycnoches pollen parent.

The awards granted to the Cycnoches ventricosum complex of species have been the most confusing. Future judging of these four species cannot rely on the awards for standards unless corrections are taken into account. I have tried to be as clear as possible in stating the differences I see among these four species. I hope the Plates I have assembled and included in this paper as well as the Key may be of assistance. As I pointed out previously, I find no fault in the quality of the plants and flowers which received awards, with the possible exception of the two Certificates of Botanical Recognition which I believe were erroneously granted. Cycnoches and its hybrids, with some exceptions, do not fit into the classic "round, full" ideal of the more popular genera and so in most instances are best judged on the General Scale.

I consider it essential that the sex of the flowers be determined before judging any Cycnoches species or hybrid. This point also applies to the true Catasetum species and hybrids. The sex of the flowers effects the size, arrangement, number of flowers and even color in some cases. Thus it is an important factor in judging any of these clones.

A. Both male and female flowers large and very similar in superficial appearance except for column structure. Lip of all sexes fleshy, more or less convex with smooth, entire margins. Column of male flowers slender and arching, much longer than wide with pollinarium conspicuously present at the tip of the column; stigma vestigial. Column of female flowers relatively short and broad compared with that of the male flowers, with the presence of two notches or "hooks" on the sides of the column tip instead of pollinarium; stigma prominent on inner surface below the tip of the column. Section Eucycnoches

b. Length of all segments over twice as great as width; flowers variously cupped with petals generally upswept more or less framing the lateral sepals and the lip. Largest flowers of the genus usually exceeding 12 cm.

c. Claw of lip blade rather long, prominently winged; column dark purple to maroon; lip callus lacking or obscure. C. loddigesii

lip; column clear green to yellowish green, prominent triangular callus at base of the lip with shallow dark green depression. C. chlorochilon

d. Short claw to lip blade; lip ivory-colored, convex, almost round with prominent dark green, almost black, depression at the base, narrow flat edge completely circling outer edge of lip.

dd. Lip almost sessile, only slightly concave; broadly oval, pinkish color with small maroon dots. Callus at base of lip in the form of two fleshy teeth, pincher-shaped. Sepals and petals olive green variously spotted maroon. Smallest flower of the section. C. haagii

cc. Sepals and petals strongly reflexed, lip with prominent claw which is generally terete, although narrow wings sometimes present on male flowers . Lip strongly convex.

d. Apex of basal callus of lip rounded, obtuse, not strongly projecting; sepals and petals light green to yellowish-green.

dd. Apex of basal callus elongated, tongue-shaped and prominently projecting from base of almost bubble-shaped lip. Sepals and petals generally very dark green. C. lehmannii

I also refer the reader to Plates I through VI for pictures of the male and female flowers of the six species of the section Eucycnoches.

A) Cycnoches chlorochilum male flower, photographed by Jim Williams, Venezuela, 1994.

B) Cycnoches chlorochilum female flower, Cycnoches chlorochilum 'Anita Bergold 'AM/AOS 86 Points, 1966

A) Cycnoches warscewiczii male flower award as C. chlorochilon 'Don Juvenal' AM/AOS 84 points 1991

B) Cycnoches warscewiczii female flower award as C. chlorochilon 'Green Jade' HCC/AOS 79 points 1971

C)Cycnoches warscewiczii (right) C. chlorochilon (left) male flowers comparison author's picture 1994

A) Cynoches ventricosum male flower from Pridgeon (ed) Illustrated Encyclopedia of Orchids 1992

A) Cycnoches lehmannii male flower Orchid Identification Center Marie Selby Gardens (OIC).

B) Cycnoches lehmannii female flower Angel Andreetta in Native Ecuadorian Orchids Dodson & Escobar 1994

A) Cycnoches loddigesii male flower C. loddigesii 'Caripe' CCM/AOS 82 points 1969

B) Cycnoches loddigesii female flower Munich Botanical Gardens Dr. Gunter Gerlach

A) Cycnoches Chloroge 'Mamasco' HCC/AOS female flower 78 points 1983 (C. chlorochilon X C. loddigesii)

B) Cycnoches Pistachio Moon 'The Prophet' HCC/AOS female flower 77 points 1981

A) Cycnoches maculatum male flower Munich Botanical Gardens Dr. Gunter Gerlach

B) Cycnoches maculatum female flower Munich Botanical Gardens Dr. Gunter Gerlach

A) Cycnoches dianae male flower, green form From The Orchid Atlas Vol 6. Karasawa 1994

B) Cycnoches dianae male flower, pink form, Award to C. egertonianum var dianae 'Woodpoint' CHM/AM/AOS 83 points 1987

A) Cycnoches peruvianum male flower Angel Andreetta from Native Ecuadorian Orchids Dodson & Escobar 1994

B) Cycnoches herrenhusanum male flower Rudolf Jenny from Die Orchidee 42(2) 1991

A) Catanoches (Clowenoches) Rebecca Northen Rebecca Northen from Orchid Review 91 (1078) 1983

B) Cycnodes Wine Delight, possibly female dominant flower Cynd. Wine Delight ' J.E.M.' FCC/AOS 91 points, 1984

NOTE: Bibliographic references to the type descriptions of species are found in the Classification
appendix attached and are not duplicated here. References marked *
indicate sources I was not able to consult directly. The information was
derived from other sources listed.

Ackerman, J.D. 1983. Society and mutual dependency of the orchid-Euglossine bee
interaction. Biological Journal of the Linnean Society 20: 301-314.

Allen, P.H. The Swan orchids A Revision of the Genus Cycnoches. Orchid Journal
1:173-184, 225-230, 249-254, 273-276, 297-303.

Ames, 0. 1944. Destruction of the Schlechter Herbarium by Bombing. A.O.S. Bulletin 13(4): 105,106.

______ 1960. The Orchidaceae of Costa Rica - Part S Cycnoches Lindl. The Orchid
Weekly 2(33): 327-328.

______ and D.S. Correll 1952. Orchids of Guatemala. Fieldiana, Botany, 26: (2)
518-522.

Batchelor, S.R 1983. Catasetum and Cycnoches A.O.S. Bulletin 52: part 1(6) 605-611
and back cover;part2 (7)713-721; part 3(8)829-835; part4
(9)917-924; part 5(10)1051-1055; part6 (11)
1163-1169; part7(12) 1295-1304.

Bechtel, H., P. Cribb and E. Launert 1986 (second edition). The Manual of Cultivated
Orchid Species. Cambridge MA: The MIT Press. Cycnoches pages 100-102, color plates 189.

Bergold, G.H. 1960. An unusual bigeneric cross. A.O.S. Bulletin 29(8): 585-588.

Blowers, J.W. 1957. The intriguing Cycnoches egertonianum. Orchid Review 65:
194-196.

Butcher, H.P. 1964. Mormodes and Allies. Orchid Weekly 5(6): 54, 55.

Capriles, AH. 1991. An Evaluation of Catasetinae Hybridizing. Orchid Digest 52(2):
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Cavestro, W. 1977. The Culture of Catasetum, Monnodes and Cycnoches. A.O.S.
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Chase, M.W. and J.S. Pippen 1990. Seed Morphology and Phylogeny in Subtribe
Catasetinae (Orchidaceae). Lindleyana 5(2): 126-133.

Chong, J.G., M.S. Strauss and J. Arditti 1982. Flower Induction and Physiology in Orchids.
Orchid Biology, Reviews and Perspectives II Chap. 7: 218-241.

Coffinan, R 1968. Propagating Cycnoches by carelessness. A. 0.5. Bulletin 34(11): 966

Cooper, E. 1929. Some Cycnoches species. Orchid Review 34: 364-367

Cope, W.C. 1942. Cycnoches chlorochilon. A.O.S. Bulletin 11(1): page unnumbered.

* Darwin, C. 1862. On the three remarkable sexual forms of Catasetum fridentatum in the possession of the Linnean Society Proc. Linn. Soc., BoL 6: 151-157.

______ 1877. The various contrivances by which orchids are fertilised by insects. Second edition,
revised. Chicago: University of Chicago Press. Chapter VII: 178-225.

Dillon, G. W. 1943. Cycnoches warscewiczii . A.O.S. Bulletin 40(1): 20 and frontispiece.

______ 1971. Orchids, Man and Nature. A. O.S. Bulletin 40(1): 39,40.

Dodson, C.H. 1962(a). Pollination and variation in the subtribe Catasetinae (Orchidaceae). Ann. Missouri Bot Gard. 49: 35-56

______ 1962). The importance of pollination in the evolution of the orchids of tropical America.
A.O.S. Bulletin 31(7): 525-534; (8) 641-649, (9) 731-735.

*_______ 1970. The role of chemical attractants in orchid pollination. Pages 83-107 in K. L. Chambers, ed. Biochemical coevolution. Oregon State University Press. Corvallis.

______ 1975. Dressleria and Clowesia: A new genus and an old one revived in the Catasetinae
(Orchidaceae). Selbyana I: 130,131.

_______ and G. P. Frymire 1959. Cycnoches lehmannii. A.O.S. Bulletin 28(3): 181-185.

*______ and _____ 1961. Natural Pollination of Orchids. Missouri Botanical Gardens Bulletin
49:133-152.

______ and H.G. Hills 1966. Gas chromatography of orchid fragrances. A.O.S Bulletin 35(9) 720 - 725.

_______and R. J. Gillespie 1967. The Biology of the Orchids. Nashville: The Mid-America Orchid Congress, Inc.

________R.L. Dressler, H.G. Hills, R M. Adams and N.H. Williams. 1969. Biologically active compounds in orchid fragrances. Science 164: 1243-1249.

_______ and P. M. Dodson 1982. Cycnoches peruviana. Icones Plantarum Tropicarum: Plate 420.

_______ and D. Bennett 1989. Cycnoches cooperi. Icones Plantarum Tropicarum Series II: Plate 0038.

_______ and _______ 1989. Cycnoches haagii. Icones Plantarum Tropicarum Series II: Plate 0039.

_______and _______ 1989. Cycnochespentadactylon. Icones Plantarum Tropicarum Series II: Plate 0041.

_______ and _______ 1989. Cycnoches peruviana. Icones Plantarum Tropicarum Series II: Plate 0042.

_______ and R Vasquez 1989. Cycnoches haagii. Icones Plantarum Tropicarum Series II: Plate 0317.

_______ and R. Escobar R 1994. Native Ecuadorian Orchids 1:164-167. Medellin: Compania Litografica Nacional S.A.

* Dressler, R.L. 1967. Why do Euglossine bees visit orchid flowers? Atas simposio sobre biota Amazonica 5 (Zool.): 171-180.

_______ 1968(a). Pollination by Euglossine bees. Evolution 22: 202-210.

*_____ 1968(b). Observations on orchids and Euglossine bees in Panama and Costa Rica. Revista de Biologia Tropical 15: 143-183.

_______1981. The Orchids Natural History and Classification. Cambridge MA: Harvard University Press.

_______1993(a). Field guide to the orchids of Costa Rica and Panama. Ithaca. Cornell University Press.

______ 1993(b). Phylogeny and Classification of the Orchid Family Portland OR: Dioscorides Press

Dunn, H..A. 1948. A swan orchid of Panama. A. O.S. Bulletin 17(6): 362

Dunsterville, G.C.K. 1967. Serendipity. OrchidReview 75(883): 18-20.

Geber, M.A. 1995. Fitness effects of sexual dimorphism in plants. Tree 10(6): 222,223.

* Gregg, K.B. 1973. Studies on the Control of Sex Expression in the General Cycnoches and Catasetum, Subtribe Catasetinae, Orchidaceae. Ph.D. Dissertation, University of Miami, Coral Gables, Fl.

_______ 1975. The effect of light intensity on sex expression in species of Cycnoches and Catasetum (Orchidaceae). Selbyana I: 101-113.

_______ 1976. The interaction of light intensity, plant size and nutrition in sex expression in Cycnoches (Orchidaceae). Selbyana II: 212-223.

_______ 1982. Sunlight ehanced ethylene evolution by developing inflorescences of Catasetum and Cycnoches and its relation to fernale flower production. Botanical Gazette 143(4): 466-475.

_______ 1983. Variation in floral fragrances and morphology: Incipient Speciation in Cycnoches? Botanical Gazette 144(4): 566-576.

Hamer, F. 1975. Las Orquideas de El Salvador I: 132-138.

_______ 1982(a). Cycnoches egertonianum. Icones Plantarum Tropicarum: Plate 645.

_______ 1982(b). Cycnoches guttulatum. Icones Plantarum Tropicarum: Plate 646.

_______ 1982(c). Cycnoches pachydactylon. Icones Plantarum Tropicarum: Plate 647.

_______ 1982(d). Cycnoches ventricosum. Icones Plantarum Tropicarum: Plate 648

_______ 1985. Cycnoches stenodactylon. Icones Plantarum Tropicarum: Plate 1212.

Hamilton, RM. 1991. Bateman and Skinner: TAB. 5 - Cycnoches ventricosum. A.O.S. Bulletin 60(5): 434-436.

Hawkes, A.D. 1950. The Major Genera of Cultivated Orchids-VI Cycnoches. A.O.S. Bulletin 19(10): 559.

_______ 1951. Notes on the Genus Cycnoches. Orchid Journal I: 34-37.

_______ 1953. A note of Cycnoches chlorochilon. Orchid Journal II: 479

_______ 1958(a). Ouestion column, Cycnoches. South Florida Orchid Weekly 1(1): 4, 5.

_______ 1958(b). A note on the culture of Cycnoches egertonianum. Orchid Weekly 1(9): 98, 99.

_______ 1959. Ouestion column, short-lived species. Orchid Weekly 1(50): 504, 505.

_______ 1961. The marvelous "swan orchids"--Genus Cycnoches. Orchid Weekly 2(47):463-467

______ 1962. New Orchid Hybrids. Orchid Weekly 3(30): 291. (Cycnodes L. Sherman Adams)

_______ 1963 (a). New Orchid Hybrids. Orchid Weekly 4(13): 129. (Cycnoches Tawny)

_______ 1963(b). Paul H. Allen Dies. Orchid Weekly 4(35): 341.

______ 1965. Encyclopaedia of Cultivated Orchids. London. Faber and Faber. Cycnoches 137-139, color plate IVa

_______ 1967.Catasetum - Cycnoches hybrid.Orchid Weekly 5(38): 375, 376

Hayes, E. 1994. The Subtribe Catasetinae. South African Orchid Journal 25(1): 2-7; 25(3): 101-109

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Horich, C.K. 1960. Roaming Costa Rica for Orchids.Orchid Weekly 2(15): 141-149.

_______ 1963.Polycycnis barbata. A.O.S. Bulletin 32(4): 295-298.

International Orchid Commission 1993. The Handbook on Orchid Nomenclature and Registration, fourth revised edition

Jenny, R 1984.Cycnoches egertonianum. Die Orchidee 35(4): four pages unnumbered.

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* Jones, K., and J.G. Daker. 1968. The chromosomes of orchids: III. Catasetinae. Kew Bull 22: 421-237.

Karasawa, K. 1994.Orchid Atlas. Vol. 6: 237; plates 123 through 137. Orchid Atlas Publishing Society.Tokyo.

Lasser, T. (ed.) 1970.Flora de VenezuelaXV(4) 76: 3648.(Cycnoches).Caracas. Instituto Botanico.

Lindley, 3.1843.Cycnoches ventricosum and Cycnoches egertonianum. Botanical Register 1843: Miscellaneous: 75-77.

Lindley, 3.1846. Cycnoches egertonianum; var. viride. Botanical Register t. 46.

Lumer, C. 1978. Pollination Ecology in the Orchidaceae (in two parts). Orchidata Jan: 7-18; April: 5-
16.

McNab, J. 1933. Cycnoches. Orchid Review 41: 23,24.

Merle, JUL. 1951. The graceful swan orchid. A. O.S. Bulletin 20(10): 616-617, and front cover.

* Moir, W.W.G. 1977. The Catasetinae. Orchidata 16(4): 5-7.

Moir, W.W.G. and M. 1959. Orchid Collecting in Panama. Orchid Weekly 1(32): 321-326.

Moimier, G. 1987. Catasetum breeding for flower longevity. Florida Orchidist 30(3): 113-121.

_______ 1989. Breeding members of Subtribe Catasetinae (Monnodes, Cycnoches, Catasetum) for
flower longevity. Orchid Digest 53(1): 4-8.

______ 1990. A current history of Catasetinae breeding. A.O.S. Bulletin 59: part 1(10) 994-998 and front cover; part 2(11) 1124-1128; part 3 (12)1221-1226.

_______ 1992. Sexual polymorphism in the genus Mormodes. Florida Orchidist 34(4): 180-184.

_______ 1994. Confusion in the Catasetinae. Orchid Digest 58(3): 143-145.

Moore, A. 1960. Cycnoches chlorochilon. Orchid Digest 24: 454, 455.

Mora, D. E. and 3. T. Atwood 1992. Cycnoches egertonianum. Icones Plantarum Tropicarum: Plate 1418.

_______ and ______ 1993. Cycnoches warscewiczii. Icones Plantarum Tropicarum: Plate 1514.

Morgan, 3. 1909. Calendar of Operations for April. Orchid Review 17: 108-111.

Northen, R.T. 1973. A glimpse into the world of species. A.O.S. Bulletin 42(1): Cycnoches 1007 and photograph 1004.

_______ 1983. Catanoches Rebecca Northen. Orchid Review 91(1078): 258.

van der Pijl, L. and C.H. Dodson 1966. Orchid Flowers: Their Pollination and Evolution. Coral Gables:University of Miami Press.

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Pring, G.H. 1943. Five-fingered swan-neck orchid (Cycnoches pentadactylon). A.O.S Bulletin 11(12):410-412.

Ramsey, C.T. 1950. The triggered rostellum of the genus Listera. A.O.S. Bulletin 19 (9): 482485.

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Roccaforte, M. and S. 1974. Cycnoches ventricosum var. warscewiczii 'Green Glow', CCM/AOS. A.O.S Bulletin 43(5): 394, 395, and front cover.

* Rolfe, R A. 1889. On the sexual forms of Catasetum with special reference to the researchers of Darwin and others. J. Linn. Soc. Bull206-225.

_______ 1893(a). Cycnoches pentadactylon. Orchid Review 1: 73-75.

______ 1893(1,). Cycnoches loddigesii. Orchid Review 1(6): 165, 166.

_______ 1895.Cycnoches warscewiczii andC. arum. Orchid Review 3(33): 263, 264.

_______ 1896.Cycnoches maculatum. Orchid Review 4(44): 228, 229.

_______ 1898. The late Mr. James Bateman. Orchid Review 6(62):56, 57

. _______ 1907(a).Cycnoches loddigesii. Orchid Review 15: 25, 26.

_______ 1907(1,).Cycnoches egertonianum. Orchid Review 15: 337-340.

_______ 1908(a). Orchids: their structure, development, and fertilisation. Orchid Review 16:215-219.

_______ 1908(1,).Cycnoches egertonianum. Orchid Review 16: 296-299

_______ 1908(c).Cycnoches maculatum. Orchid Review 16: 370, 371

_______ 1909(a).Cycnoches densiflorum. Orchid Review 17: 104, 105.

_______ 1909(1,). Seeds of Cycnoches chlorochilon. Orchid Review 17: 168

_______ 1909(c). Cycnoches maculatum. Orchid Review 17: 273, 274.

_______ 1909(4).A revision of the genus Cycnoches. Bulletin of Misc. Information ( Kew Bulletin)
XXXIV: 268-277.

_______ 1910(a). Cycnoches stelliferum and egertonianum. Orchid Review 18: 253.

_______ 1910(1,). The evolution of the Orchidaceae. Orchid Review 18(215): 321-325.

_______ 1911. Cycnoches stelliferum. Orchid Review 19: 255, 256.

_______ 1912. Cycnoches chlorochilon: Female flower. Orchid Review 20: 329, 330.

_______ 1917. Mistakes in orchidology. Orchid Review 25: 150-184.

_______ 1918. Cycnoches ventricosum. Orchid Review 26: 189-191.

_______ 1918. Cycnoches chlorochilon. Orchid Review 26: 209, 210.

_______ 1919(a). Cycnoches stelliferum. Orchid Review 27: 162.

_______ 1919(1,). Cycnoches ventricosum from Chiriqui. Orchid Review 27: 175, 176.

_______ 1919(c). Cycnoches dianae. Orchid Review 27: 107, 108.

Romero, G.A. 1990. Phylogenetic relationships in subtribe Catasetinae (Orchidaceae, Cymbidieae).
Lindleyana 5(3): 160-181.

______ and R Jenny 1993. Contributions toward a monograph of Catasetum Catasetinae, Orchidaceae
I:A checklist of species, varieties, and natural hybrids. Harvard Papers 4: 59-84.
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Ltd. Cycnoches: pp.134-136.

Schlechter, R 1916. Die Gattung Cycnoches Ldl. Orchis 10: 47-61. Personal translation from German into English 1994 by Dr. Ursula Hoffman for this paper, The Genus Cycnoches Lindl.

Scully, R M. 1951. Growing the unusual orchids. A.O.S. Bulletin 20(4): 209 Cycnoches chlorochilon

Segars, M. 1963. Conversation piece--the curious Cycnoches. Orchid Digest 27(5): 196-198.

Segars, W. E. 1960. Notes on the Culture of Cycnoches. A. 0. S Bulletin 29(8): 582-584.

______ 1962. Orchid Cultural Calendar for November-What to do in Florida. A. 0. S Bulletin 31(11): 890.

_______ 1963 (a). Notes on the culture of the Catasetinae. A.O.S Bulletin 32(4): 268-270.

______ 1963(1,). New Hybrid of Cycnoches. A. 0. S Bulletin 32(4): 293, 294.

_______ 1963(c). Notes on the culture of the Catasetinae. Orchid Digest 27(5): 198, 199.

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_______ 1994.An Illustrated Survey of Orchid Genera, Cycnoches: pp.130, 131.

Slocum, C.P. 1961. Cycnodes Sherman Adams: first of a new genus. A. O.S. Bulletin 30(9): 728.

Teuseher, H. 1955. Cycnoches aureum. A.O.S. Bulletin 24(3): 171, 172.

_______ 1961. Four little-known species of the genus Mormodes. A.O.S. Bulletin 30(8): 648-655.

_______ 1962. Cycnoches cooperi. A.O.S. Bulletin 31(7): 524, 556, 557.

_______ 1963. Cycnoches cooperi. A.O.S. Bulletin 32(8): 622-624.

_______ 1964. Cycnoches pentadactylon. A. O.S. Bulletin 33(1): 20-22.

_______ 1965(a). Cycnoches egertonianum. A. O.S. Bulletin 34(5): 393-396.

_______ 1965). A novel method of growing Catasetum. A.O.S. Bulletin 34(12): 1067, 1068.

Unidentified Author 1900. Cycnoches pentadactylon. Orchid Review 8: 312, 313.

______ 1910. Cycnoches at Kew. Orchid Review 18: 315.

_______ 1912(a). Orchids at Kew. Orchid Review 20:315, 316.

_______ 1912(b). Cycnoches densiflorum. Orchid Review 20: 331.

______ 1917. Variations of Hybrid Orchids. Orchid Review 25: 53.

_______ 1922. Cycnoches loddigesii. Orchid Review 30: 139, 140.

_______ 1931. Cycnoches pentadactylon. Orchid Review 39: 368.

_______ 1932. Cycnoches chlorochilon. Orchid Review 40: 240, 241.

_______ 1943. Cycnoches warscewicrii. A.O.S. Bulletin 12(1): frontispiece and 20.

______ 1966. Notes from Panama: Cycnoches chlorochilon. Orchid Review 74(871): 23.

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Vandeae
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Wells, E. W. 1950. Orchids and gardens of the Southern States and Caribbean Gulf Area 11. A.O.S.
Bulletin 19(10): 544

Werkhoven, M. C. M. 1986. Orchids of Suriname. Cycnoches: 110.

Williams, B.S. and H. 1894. Cycnoches pp.215-217 in The Orchid Grower's Manual seventh edition

Victoria & Paradise Nurseries. London. Reprint 1961, Weldon & Westley Ltd. Codicote,
Herts.

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Williams, N.H. 1982. The biology of orchids and Euglossine bees. pp.119-171 in Orchid biology reviews
and perspectives. Vol.2, J. Arditti, ed. Cornell University Press, Ithaca, NY

Williams, N.H. and C.FL Dodson 1972. Selective attraction of male Euglossine bees to orchid
floral fragrances and its importance in long distance pollen flow. Evolution 26(1): 84-95.

APPENDIX A - CLASSIFICATION Prepared by G. Carr
Based on Phylogeny and Classification of the Orchid Family by R. L. Dressler, published 1993.
FAMILY: Orchidaceae
SUBFAMILY: Epidendroideae
PHYLAD: Cymbidioid
TRIBE: Cymbidieae
SUBTRIBE: Catasetinae
GENUS: Cycnoches Lindley 1832 The Genera and Species of Orchidaceous Plants.
SPECIES: --listed by Section in order of date of description and publication.
To my knowledge, no monograph of the genus Cycnoches has ever been published. Three revisions, Rolfe 1909, Schlechter 1916 and Allen 1952, have been published. They form the basis for this list, updated and modified with information from Dr. K. Senghas (1995 personal communication), supported by the works of Dr. C. Dodson, Dr. R. Dressler, Dr. K. Gregg, Dr. Gunther Gerlach, Dr. Gustavo Romero, Dr. Rudolf Jenny and the sources listed below for the description of new species since 1989. Additional references to species are found in the various Icones Planatarum Tropicarum series published first by the Missouri Botanical Gardens and currently by Selby Gardens, and the contribution by Dr Senghas on the genus published in the Third Revised edition of Schlechter's Die Orchideen, Vol. IB, fascicle 26, 1602-1608. These references are found in the Bibliography under the authors' names. Any errors that might occur are my responsibility and not that of any of my sources. This listing is offered as a guide to what I have found in my research.

Species marked * have been placed in synonymy with the species preceding their listing instead of the normal usage of = to signify synonymy. With the exception of Allen's varieties (1952), I have omitted the listing for species which were reduced to varieties at one time, but which are considered here as valid species. For example, since I am listing C. cooperi Rolfe as a valid species, I have omitted the reference to it as C. pentadactylon var. cooperi Pabst.

EUCYCNOCHES Rolfe 1909

1 loddigesii Lindley, 1832 Gen. & Sp. Orch. Pl. 154.
var. leucochilum Hooker 1841 Bot. Mag. t. 3855.
var. purpureum hort. ex Reichb. f. 1857 Cat. Orch.-Samml. Schiller 22
*cucullata Lindley 1837 Bot. Reg. 10: t. 1951.
2 chlorochilon Klotzsch 1838 Allgem. Gartenz. 6: 225.
* ventricosum var. chlorochilon Allen 1952 Orchid Journal I: 297 ff.
3 ventricosum Bateman 1842 Orch. Mex. & Guat. t. 5.
4 warscewiczii Reichenbach filius 1852 Bot. Zeit. 10: 734.
* tonduzii Schlechter 1923 Repertorium Spec. Nov. Regni Veg. Beihefte 19: 298
. * ventricosum var. warscewiczii Allen 1952 Orchid Journal I: 297 ff.
5 lehmannii Reichenbach filius 1878 Otia Botanica Hamburgensis 1: 10.
6 haagii Barbosa Rodrigues 1882 Orch. Nov. 2: 221
* versicolor Reichenbach filius 1888 Gardeners' Chronicle 2: 596.
* haagei (Barb. Rodr.) Schlechter 1916 Orchis 10:55

SECTION HETERANTHAE Rolfe 1909

7 maculatum Lindley 1840 Bot. Reg. misc. 10.
8 egertonianum Bateman 1842 Orch. Mex. & Guat. t. 40.
var. viride Lindley 1846 Botanical Register 32: 1846, t. 46
*ventricosum var. egertonianum Hooker 1844 Bot. Mag. t. 4054.
*stelliferum Loddiges ex Reichenbach f. 1863 Ann. Bot. Syst. 6: 561
* rossianum Rolfe 1891 Gardeners' Chronicle 1: 456
*glanduliferum Rolfe 1892 Gard. & For. 5: 90.
*amparoanum Schlechter 1923 Repertorium Spec. Nov. Regni Veg. Beihefte 19: 48.
* pauciflorum Schlechter 1923 Repertorium Spec. Nov. Regni Veg. Beihefte 19: 137.
9 pendadactylon Lindley 1843 Botanical Register, misc. 18, t. 22.
*amesianum Sander 1902 Gardeners' Chronicle 1: 218.
*espiritosantense Brade ex Hoehne 1942 Fl. Brasil. 12 (6): 145, t 105.
10 aureum Lindley 1852 Paxton's Flower Garden 3: 6, t. 75.
egertonianum var. aureum Allen 1952 Orchid Journal I: 297 ff.
11 dianae Reichenbach filius 1852 Bot. Zeit. 10: 636.
* albidum Kraenzlin 1909 Repertorium Spec. Nov. Regni Veg. 7: 38.
*powellii Schlechter 1922 Repertorium Spec. Nov. Regni Veg. Beihefte 17: 58.
*egertonianum var. dianae Allen 1952 Orchid Journal I: 297 ff.

12 peruviana Rolfe 1891 Lindenia 7: t. 301.

13 densiflorum Rolfe 1909 Kew Bull. 1909: 63
14 cooperi Rolfe 1913 Kew Bull. 1913: 143
15 guttulatum Schlechter 1922 Repertorium Spec. Nov. Regni Veg. Beihefte 17: 56.
16 pachydactylon Schlechter 1922 Repertorium Spec. Nov. Regni Veg. Beihefte 17: 57.
17 stenodactylon Schlechter 1922 Repertorium Spec. Nov. Regni Veg. Beihefte 17: 59.
18 brachydactylon Schlechter 1924 Repertorium Spec. Nov. Regni Veg. Beihefte 27: 80

19 bennettii Dodson 1989 Icones Plantarum Tropicarum, Series II Plate 0037
20 jarae Dodson 1989 Icones Plantarum Tropicarum, Series II Plate 0040
21 suarezii Dodson 1989 Icones Plantarum Tropicarum, Series II Plate 0431
22 thurstonorum Dodson 1989 Icones Plantarum Tropicarum, Series II Plate 0432
23 herrenhusanum Jenny & Romero 1991 Die Orchidee 1991: 42(2) 68-72.
24 quatuorchristis Bennett 1992 Lindleyana 7(2) 83-85, 87
25 manoelae Castro & Campacci 1993 Boletim Circulo Paulista de Orquidofilos V (1): 25-26

CYCNOCHES - NOMEN NUDUM I found references to two species, for which I cannot locate a valid description. They are listed here with source.

Cycnoches lehmanniana - Name is found on herbarium sheets in the Reichenbach herbarium, but there was never a valid publication of the name.
As best as I can see, the shape of the lip and the long claw would indicate it is really Cycnoches lehmannii.
Cycnoches Forgetii - Listed in Sanders' Orchid Guide, 1927 edition on page 135 with the notation that it is similar to C. peruvianum and C. cooperi.
The place of origin is given as Peru, and I assume it was collected by the French collector Forget who collected in Peru in the early
1920's. Without details, it is impossible to identify, but presumably, it must have been identified as one of the two species to which the Guide referred.

SPECIES ORIGINALLY DESCRIBED AS MEMBER OF THE GENUS CYCNOCHES BUT WHICH HAVE BEEN TRANSFERRED TO OTHER GENERA
Cyc. barbatum Lindley 1849 Journal of the Horticultural Society 4: 268 = Polycycnis barbata
Cyc. pescatorei Lindley 1850 Paxton's Flower Garden 1: 123. = Lueddemannia pescatorei
Cyc. musciferum Lindley 1852 Paxton's Flower Garden 3:28, fig. 248. = Polycycnis musciferia
Cyc. viride Koch 1857 Berl. Allgem. Gartenz. 25: 291 = Clowesia russelliana
Cyc. lindleyi hort ex Linden 1860 Pescat. sub t. 22. = Lueddemannia pescatorei
Cyc. buchtieni Kraenzlin 1928 Repert. Spec. Nov. Regni Veg. 25: 26. = Polycycnis buchtieni

APPENDIX B - HYBRIDS REGISTERED THROUGH OCTOBER 1997

CYCNOCHES Lindley

NAME POD PARENT POLLEN PARENT YEAR REGISTRANT (HYBRIDIZER)

Hawaiian Lei egertonianum * dianae 1961 W.W.G. Moir

Tawny egertonianum chlorochilon ** 1962 Rivermont Orchids

Chloroge chlorochilon ** loddigesi 1970 Polk/Mamasco

Cygnet chlorochilon ** haagii 1970 H. L. Allen
Rip-Van-Winkle maculatum chlorochilon ** 1975 W. Osment

Tres Colores chlorochilon ** Tawny 1976 A. M. Schmidt

Pistachio Moon egertonianum haagii 1978 Jones & Scully

Nova Luna Pistachio Moon loddigesii 1984 Mrs. R. Levy

Bo Day Shus haagii loddigesii 1987 J.E.M. Orchids

Pentalodd pentadactylon loddigesii 1991 J.E.M. Orchids

Rocky Clough pentadactylon egertonianum 1995 G. Carr (H&R Nurseries)

Pentawar warscewiczii pentadactylon 1996 G. Carr (Carter & Holmes)

Jumbo Dragon Chloroge 'Golden Sand' pentadactylon 1996 Amir Trading / Jumbo Orchids

Mass Confusion chlorochilon warscewiczii 1997 G. Carr (C. Clouse)
* According to Jenny et al 1995, the pod parent was C. aureum which the Registration Authority considers synonymous with C. egertonianum ** As explained in Part I of this paper, it is probable that the actual species used in these crosses was C. warscewiczii.
CYCNODES = Cycnoches Lindl. X Mormodes Lindl.

NAME POD PARENT POLLEN PARENT YEAR REGISTRANT (HYBRIDIZER)

L. Sherman Adams C. chlorochilon M. wendlandii 1961 Jones & Scully / Slocum

Ginger Snap C. chlorochilon M. colossus 1966 Alberts & Merkel Bros.

Thomas Krumpe M. buccinator C. chlorochilon 1969 Forman & Krumpe

Wine Delight C. lehmanni M. sinuata 1980 Coffman (Fuchs Jr.)

Chocolate and Cherries C. chlorochilon M. sinuata 1987 J.E.M. Orchids

Key Lime Pie C. haagii M. buccinator 1990 G. Monnier

Raspberry Yogurt M. frymirei C. pentadactylon 1991 J.E.M. Orchids

Jumbo Phoenix C. Chloroge 'Jumbo' M. sinuata 'Ruby' 1996 Amir Trading / Jumbo Orchids

CATANOCHES *** = Catasetum Rich. ex Kunth X Cycnoches Lindl.

NAME POD PARENT POLLEN PARENT YEAR REGISTRANT (HYBRIDIZER)

Crazy Creature Cyc. chlorochilon Ctsm. Orchidglade 1978 Jones & Scully

Fantasy Ctsm. saccatum Cyc. chlorochilon 1978 Rod McLellan

Lemon Meringue Cyc. haagii Ctsm. Orchidglade 1982 Jones & Scully

Penthouse Cyc. pentadactylon Ctsm. tenebrosum 1982 Jones & Scully

Green Dragon Cyc. haagii Ctsm. expansum 1986 J.E.M. Orchids

Midnight Jem Cyc. chlorochilon Ctsm. Susan Fuchs 1991 J.E.M. Orchids

Yellow Bird Cyc. chlorochilon Ctsm. pileatum 1991 J.E.M. Orchids

Brown Sugar Cyc. egertonianum Ctsm. callosum 1996 G. Carr (Charles Alford)

Bela Vista Cyc. haagii Ctsm. denticulatum 1996 Antonio Schmidt

Alvaro Pereira Ctsm. pileatum Cyc. pentadactylon 1997 L. Pereira dos Santos
(CLOWENOCHES *** = Clowesia Lindl. X Cycnoches Lindl.) .

Green Beret Clo. warscewiczii Cyc. ventricosum 1967 W.W.G. Moir

Rebecca Northen Clo. rosea Cyc. chlorochilon 1973 Furrow & Co., Inc
Green God Cyc. chlorochilon Clo. warscewiczii 1980 W. Osment

Lunar Cheese Cyc. haagii Clo. russelliana 1982 Jones & Scully

*** In 1975, C. H. Dodson undertook to simplify the genus Catasetum by reestablishing the genus Clowesia Lindley and erecting a new genus Dressleria Dodson. The species which had been included in Catasetum Rich. ex Kunth, which have perfect (hermaphroditic) flowers were removed to these two genera as appropriate. This left the genus Catasetum to include only those species with dimorphic flowers. In line with this taxonomic revision, new intergeneric names have been proposed for hybrids involving what are now considered to be species of Clowesia and Dressleria. The Registration Authority however, has chosen not to recognize this taxonomic change. Therefore, any hybrid which involves species now considered to be Clowesia or Dressleria are still registered as hybrids of Catasetum. Since Dodson's revision, new species have been described and published which have been directly placed into the genera Clowesia and Dressleria. When these new species are eventually used in hybridization, it should be interesting to see if the Registration Authority will then accept Dodson's revision. The first usage of Clowenoches I found was in an article by Rebecca Northen written for the Orchid Review (Northen 1983). In a footnote in discussing "her" Catanoches, Ms. Northen suggested that perhaps the hybrid should be more correctly called Clowenoches. In an article in L'Orchidophile, a complete set of names for intergeneric hybrids within the Catasetiinae was proposed - see Jenny et al 1995. For the sake of clarity, I have chosen to follow this proposal and have divided the registered Catanoches into two parts.
CYCNANDRA (CYCDA.) = Galeandra Lindl. X Cycnoches Lindl.

NAME POD PARENT POLLEN PARENT YEAR REGISTRANT (HYBRIDIZER)

Jumbo Pioneer G. devoniana 'Solar Wind' C. haagii 'Green Gold' 1996 Amir Marketing / Jumbo Orchids

Jumbo Spirit G. devoniana 'Solar Wind' C. pentadactylon 'Galaxy' 1996 Amir Marketing / Jumbo Orchids

MONNIERARA = Cycnoches X Mormodes X Catasetum

NAME POD PARENT POLLEN PARENT YEAR REGISTRANT (HYBRIDIZER)

Ruby Matrix Cycd. Wine Delight Ctsm. expansum 1990 J.E.M. Orchids

Mary Rose Cycd. Wine Delight Ctsm. Susan Fuchs 1991 J.E.M. Orchids

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Plate I

Plate II

Plate III

Plate IV

Plate V

Plate VI

Plate VII

Plate VIII

Plate IX

Plate X

Plate XI

Plate XII

Plate XIII

Plate XIV

Plate XV

BIBLIOGRAPHY

NAME	POD PARENT	POLLEN PARENT	YEAR	REGISTRANT (HYBRIDIZER)
Hawaiian Lei	egertonianum *	dianae	1961	W.W.G. Moir
Tawny	egertonianum	chlorochilon **	1962	Rivermont Orchids
Chloroge	chlorochilon **	loddigesi	1970	Polk/Mamasco
Cygnet	chlorochilon **	haagii	1970	H. L. Allen
Rip-Van-Winkle	maculatum	chlorochilon **	1975	W. Osment
Tres Colores	chlorochilon **	Tawny	1976	A. M. Schmidt
Pistachio Moon	egertonianum	haagii	1978	Jones & Scully
Nova Luna	Pistachio Moon	loddigesii	1984	Mrs. R. Levy
Bo Day Shus	haagii	loddigesii	1987	J.E.M. Orchids
Pentalodd	pentadactylon	loddigesii	1991	J.E.M. Orchids
Rocky Clough	pentadactylon	egertonianum	1995	G. Carr (H&R Nurseries)
Pentawar	warscewiczii	pentadactylon	1996	G. Carr (Carter & Holmes)
Jumbo Dragon	Chloroge 'Golden Sand'	pentadactylon	1996	Amir Trading / Jumbo Orchids
Mass Confusion	chlorochilon	warscewiczii	1997	G. Carr (C. Clouse)

NAME	POD PARENT	POLLEN PARENT	YEAR	REGISTRANT (HYBRIDIZER)
L. Sherman Adams	C. chlorochilon	M. wendlandii	1961	Jones & Scully / Slocum
Ginger Snap	C. chlorochilon	M. colossus	1966	Alberts & Merkel Bros.
Thomas Krumpe	M. buccinator	C. chlorochilon	1969	Forman & Krumpe
Wine Delight	C. lehmanni	M. sinuata	1980	Coffman (Fuchs Jr.)
Chocolate and Cherries	C. chlorochilon	M. sinuata	1987	J.E.M. Orchids
Key Lime Pie	C. haagii	M. buccinator	1990	G. Monnier
Raspberry Yogurt	M. frymirei	C. pentadactylon	1991	J.E.M. Orchids
Jumbo Phoenix	C. Chloroge 'Jumbo'	M. sinuata 'Ruby'	1996	Amir Trading / Jumbo Orchids

NAME	POD PARENT	POLLEN PARENT	YEAR	REGISTRANT (HYBRIDIZER)
Crazy Creature	Cyc. chlorochilon	Ctsm. Orchidglade	1978	Jones & Scully
Fantasy	Ctsm. saccatum	Cyc. chlorochilon	1978	Rod McLellan
Lemon Meringue	Cyc. haagii	Ctsm. Orchidglade	1982	Jones & Scully
Penthouse	Cyc. pentadactylon	Ctsm. tenebrosum	1982	Jones & Scully
Green Dragon	Cyc. haagii	Ctsm. expansum	1986	J.E.M. Orchids
Midnight Jem	Cyc. chlorochilon	Ctsm. Susan Fuchs	1991	J.E.M. Orchids
Yellow Bird	Cyc. chlorochilon	Ctsm. pileatum	1991	J.E.M. Orchids
Brown Sugar	Cyc. egertonianum	Ctsm. callosum	1996	G. Carr (Charles Alford)
Bela Vista	Cyc. haagii	Ctsm. denticulatum	1996	Antonio Schmidt
Alvaro Pereira	Ctsm. pileatum	Cyc. pentadactylon	1997	L. Pereira dos Santos

Green Beret	Clo. warscewiczii	Cyc. ventricosum	1967	W.W.G. Moir
Rebecca Northen	Clo. rosea	Cyc. chlorochilon	1973	Furrow & Co., Inc
Green God	Cyc. chlorochilon	Clo. warscewiczii	1980	W. Osment
Lunar Cheese	Cyc. haagii	Clo. russelliana	1982	Jones & Scully

NAME	POD PARENT	POLLEN PARENT	YEAR	REGISTRANT (HYBRIDIZER)
Jumbo Pioneer	G. devoniana 'Solar Wind'	C. haagii 'Green Gold'	1996	Amir Marketing / Jumbo Orchids
Jumbo Spirit	G. devoniana 'Solar Wind'	C. pentadactylon 'Galaxy'	1996	Amir Marketing / Jumbo Orchids

NAME	POD PARENT	POLLEN PARENT	YEAR	REGISTRANT (HYBRIDIZER)
Ruby Matrix	Cycd. Wine Delight	Ctsm. expansum	1990	J.E.M. Orchids
Mary Rose	Cycd. Wine Delight	Ctsm. Susan Fuchs	1991	J.E.M. Orchids