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Copyright © Dr. Guido J Braem 2014
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ISBN: 9788793044012
Cover Illustration: Paphiopedilum sanderianum from the Reichenbachia. Watercolour painting by Walter Hood Fitch (1817-1892)
PREAMBLE
The deed is done. The book is completed. And we did it to the best of our ability. We present herewith the most comprehensive scientific treatise of Asian tropical slipper orchids possible, completed by nearly 400 illustrations.
In view of the ongoing discussion about the definition of species and the delineation of the various taxa, the task was a difficult one, but we think we have reached an acceptable compromise. Again there is room left for discussion, and as no book has ever been published without any errors, some will most certainly be discovered in our work. Our senior author, Prof. Dr. Braem, takes full responsibility for and claims the copyright to all typos and other calamities that may be present.
We have attempted to present a book that is attractive to all enthusiasts of the genus Paphiopedilum, amateur as well as professional growers, and to all those that just enjoy well-illustrated books on orchidaceous plants. After all, these “slippers”, as we call them, are among the most beautiful that the orchid family has to offer. Unfortunately, present legislative measures make it difficult to obtain a satisfactory amount of material of all species, in turn making scientific work on these plants rather difficult. If and when (dreams must be allowed) politicians regain their senses and do away with the misdirected law generally referred to as “CITES” as it is applied to plants and protect the habitats of the orchids, more detailed work might become possible. In the meantime, we hope that our work will inspire both professional and amateur breeders to concentrate on the in vitro propagation of these beautiful plants.
No book can be completed without help, and the authors are indebted to Alister Adhikari, P.T. Anh, Tom Ballinger, Nicolas Bougourd (La cour des orchidées), Chu Xuan Canh, Hadley Cash (Marriott Orchids), S. Chew, Dr. Jay Ciezki, Theanh Dang, Jason Fischer (Orchids Limited), Jerry Fischer (Orchids Limited), Franz Fuchs, Lourens Grobler (AfriOrchids), Gore Orchid Conservatory, Olaf Gruß, Dr. Panjai Kumar, Lan Rung Hua Binh, Prof. Dr. Zhongjian Liu, Peter Maxwell, Werner Mayer, MyOrchidsRu, Dr. Henry Oakeley, Jason Ong, Michael Ooi, Ron Parsons, Dr. Holger Perner, Linda Welsh Petchnick, Alexej Popow, Nebojsha Popow, Udai Pradhan, Alex Sánchez Sans, Eric Sauer, Sompong Sri, Montri Thanadkha, The Orchid House, Tropical Exotic Orchids, Sam Tsui (Orchid Inn Ltd.), John Varigos, Rogier van Vugt, Thang Vu, and Thanh Binh Vuong for their generous assistance. If, by oversight, we have forgotten to list someone, we apologise.
Last, but most certainly not least, we wish to thank Mr. Lars Pedersen of Moorland eBooks for his always professional and friendly cooperation.
CONTENTS
PREAMBLE
INTRODUCTION
GENERAL SECTION
Introduction to the Taxonomy and Nomenclature of the Genus Paphiopedilum
Problems in Botanical Taxonomy
The Species Concept
Training
Misidentifications
Bad Typification
Synonyms
Orchidaceae aka Orchids
General Considerations
The Orchids
The Cypripediae
The Genus Paphiopedilum
General Considerations
Morphology
Leaves
Stems and Stolons
Inflorescence
Flower Characteristics
Sepals
Petals
Lip (Labellum)
Malformations of Flower Segments
Staminode
Seeds
Ecology
Pollination
Paphiopedilum Culture (Eric R. Sauer)
Temperature
Light
Water and Fertilizer
Air Movement
Humidity
Growing Media and Repotting
Insect and Disease Issues
DESCRIPTIVE SECTION
Description of Taxa
Introduction and Conventions Used
Format of Species Description
Correct Orthography of the Taxa
Spelling of the Names of Authors
Designation with Full Citation of the Pertinent Literature
Basionym
Synonym(s)
Discussion
Etymology
Description
Distribution and Habitat
Flowering
Miscellaneous Notes
Notes on Albinism
Infrageneric Taxonomy
Preliminary Notes
Subgenus Parvisepalum
Subgenus Brachypetalum
Subgenus Paphiopedilum
Section Paphiopedilum
Section Ceratopetalum
Section Stictopetalum
Section Thiopetalum
Subgenus Sigmatopetalum
Section Sigmatopetalum
Section Spathopetalum
Subsection Macronodium
Subsection Spathopetalum
Section Blepharopetalum
Section Punctatum
Section Planipetalum
Section Barbata
Subsection Barbata
Subsection Loripetalum
Subsection Chloroneura
Subgenus Megastaminodium
Subgenus Polyantha
Section Polyantha
Section Mystropetalum
Section Mastigopetalum
Subgenus Cochlopetalum
THE SPECIES
Subgenus Parvisepalum
Paphiopedilum armeniacum
Paphiopedilum delenatii
Paphiopedilum emersonii
Paphiopedilum hangianum
Paphiopedilum malipoense
Paphiopedilum micranthum
Paphiopedilum vietnamense
Subgenus Brachypetalum
Paphiopedilum bellatulum
Paphiopedilum concolor
Paphiopedilum godefroyae
Paphiopedilum niveum
Paphiopedilum thaianum
Subgenus Paphiopedilum
Paphiopedilum charlesworthii
Paphiopedilum henryanum
Paphiopedilum herrmannii
Paphiopedilum insigne Complex
Paphiopedilum insigne
Paphiopedilum barbigerum
Paphiopedilum coccineum
Paphiopedilum exul
Paphiopedilum helenae
Paphiopedilum markianum
Paphiopedilum tranlienianum
Paphiopedilum villosum Complex
Paphiopedilum villosum
Paphiopedilum gratrixianum
Paphiopedilum fairrieanum
Paphiopedilum hirsutissimum Complex
Paphiopedilum hirsutissimum
Paphiopedilum esquirolei
Paphiopedilum druryi
Paphiopedilum spicerianum
Subgenus Sigmatopetalum
Paphiopedilum venustum
Paphiopedilum hookerae Complex
Paphiopedilum hookerae
Paphiopedilum volonteanum
Paphiopedilum sangii
Paphiopedilum appletonianum
Paphiopedilum bullenianum
Paphiopedilum cerveranum
Paphiopedilum mastersianum
Paphiopedilum mohrianum
Paphiopedilum papuanum
Paphiopedilum violascens Complex
Paphiopedilum violascens
Paphiopedilum bougainvilleanum
Paphiopedilum wentworthianum
Paphiopedilum tonsum Complex
Paphiopedilum tonsum
Paphiopedilum braemii
Paphiopedilum purpuratum
Paphiopedilum sukhakulii
Paphiopedilum wardii
Paphiopedilum argus
Paphiopedilum barbatum
Paphiopedilum crossii
Paphiopedilum fowliei
Paphiopedilum hennisianum
Paphiopedilum lawrenceanum
Paphiopedilum dayanum
Paphiopedilum superbiens Complex
Paphiopedilum ciliolare
Paphiopedilum superbiens
Paphiopedilum acmodontum
Paphiopedilum javanicum
Paphiopedilum sugiyamanum
Paphiopedilum robinsonianum
Paphiopedilum inamorii
Paphiopedilum schoseri
Paphiopedilum urbanianum
Subgenus Megastaminodium
Paphiopedilum canhii
Subgenus Polyantha
Paphiopedilum haynaldianum
Paphiopedilum lowii Complex
Paphiopedilum lowii
Paphiopedilum lynniae
Paphiopedilum richardianum
Paphiopedilum parishii Complex
Paphiopedilum parishii
Paphiopedilum dianthum
Paphiopedilum adductum Complex
Paphiopedilum adductum
Paphiopedilum anitum
Paphiopedilum elliottianum
Paphiopedilum gigantifolium
Paphiopedilum intaniae
Paphiopedilum kolopakingii
Paphiopedilum ooii
Paphiopedilum philippinense Complex
Paphiopedilum philippinense
Paphiopedilum roebbelenii
Paphiopedilum glanduliferum / praestans Complex
Paphiopedilum glanduliferum
Paphiopedilum praestans
Paphiopedilum wilhelminiae
Paphiopedilum randsii
Paphiopedilum rothschildianum
Paphiopedilum sanderianum
Paphiopedilum stonei
Paphiopedilum platyphyllum
Paphiopedilum supardii
Subgenus Cochlopetalum
Paphiopedilum chamberlainianum
Paphiopedilum kalinae
Paphiopedilum liemianum
Paphiopedilum primulinum
Paphiopedilum glaucophyllum
Paphiopedilum moquetteanum
Paphiopedilum victoria-mariae
Appendix # 1: Intrageneric Identification Keys
Appendix # 2: Index of Plant Names
Appendix # 3: Literature
Appendix # 4: List of Illustrations
INTRODUCTION
Orchids are one of the most successful groups in the plant kingdom, having conquered a niche in nearly every ecological system on earth, with the exception of the Polar Regions. They can be found in the temperate and in sub-tropical and tropical areas, in wet rainforests as well as in semi-arid and arid deserts. Whereas tropical orchids have been known to be of interest in China for hundreds of years, they reached Europe only in modern times. We know that tropical orchids were cultivated in England around the middle of the eighteenth century. However, there may well have been earlier “imports” of which there are no surviving records. We do not know when these tropical plants were first deliberately grown in the gardens of North America, but we doubt that it was much later than in Europe. Today, orchids are among the most popular groups of flowering plants all over the world.
Slipper orchids were known before 1753, the year in which Carolus Linnaeus (1707-1778) published his famous Species plantarum, the book that was the origin of the science of plant taxonomy and systematics. We find a description of Cypripedium calceolus as Calceolaria by Heister in 1748. Paphiopedilum purpuratum was in cultivation (as Cypripedium sinicum) at Loddiges’s nursery at Hackney in the British Isles before 1760. The exotic flowers of the plants we now classify under the genus Paphiopedilum have attracted interest from botanists and amateur growers since that time. By now, the genus Paphiopedilum is one of the most widespread orchid genera in cultivation. Their flowers are a unique development of nature. They are complex and interesting, and the many species show considerable variation. This variation has resulted in a large group of plants in which each and every individual is intriguing. It is, therefore, understandable why some orchid enthusiasts grow only these orchids. Furthermore, one should remember that plants of this genus were among the first to be used for artificial hybridising, the first recorded hybrid being Paphiopedilum Harrisianum1 (barbatum x villosum) made by Veitch & Sons and registered in 1869.
The literature about plants now treated under the genus Paphiopedilum was sporadic until 1894 and resulted mainly out of the discovery and subesequent description of new species. During that time all the species were treated under the genus Cypripedium (as established by Linnaeus in 1753). Some pertinent literature, such as the Flora Telluriana, published by Rafinesque in 1837 and 1838, was overlooked or not considered important. Although Paphiopedilum was established as an autonomous genus by Pfitzer in 1886, no extensive study was published before 1894. Between 1894 and 1903 the literature about Paphiopedilum and slipper orchids in general was plentiful. After 1903 it ceased abruptly. There was really no complete treatment of the genus Paphiopedilum between 1903 and 1987, and literature about Paphiopedilum remained scarce until, in the early nineteen seventies, the late Dr. Jack Fowlie (1929-1993) took an interest in the genus. Jack, a medical doctor by profession, travelled widely and was a prolific author. In his function as editor of the Orchid Digest, Fowlie published a series of articles on the genus Paphiopedilum, articles that have by now become true classics, notwithstanding the fact that his understanding of taxonomy and systematics did not always stand up to scientific scrutiny.
Since Fowlie, many articles on the genus have been published. The great majority of these have provided more or less specific information on a single species, a group of related species, or the species occurring in a defined geographical area. Several books were published, most of which were not very helpful in respect to the clarification of the natural history of the genus. Near the end of the nineteen eighties, P. J. Cribb (1987) and Guido Braem (1988) each published a monograph on the genus. These two monographs soon became the standard works in respect to Paphiopedilum. However, neither book included much on the culture of the plants. As some species were described shortly after the appearance of Cribb’s work, Braem’s monograph was, at that time, the most complete treatment of the genus.
Since the publication of Braem’s book in 1988, many new species have been discovered and described, and the information about many other species has been updated.
Interest in the genus Paphiopedilum is as high as it has ever been. Restrictive, and in some instances irrational, legislation has made the collecting and the commercial trade of these magnificent plants difficult, sometimes (or in some countries) even impossible. This in turn has raised the demand for these plants, thus stimulating the search for new species. In addition, the political changes which have occurred on our planet over the last forty years have made it possible to visit regions which have been off-limits to collectors for a long time, and this has resulted in the discovery of new species and varieties.
By 1998 Cribb had published a revision of his Paphiopedilum monograph and had contributed to some regional floras. Thus, there was need for a new, more extensive and updated monograph of the genus. In response to this need, Braem, together with Charles and Margaret Baker, revised his earlier monograph. The project was scheduled to be published in three volumes, the first of which appeared in 1998, the second in 1999. Unfortunately, due to circumstances beyond the control of the authors, the project had to be abandoned before the publication of the third volume. Complaints regarding these unfortunate proceedings are generally addressed to the authors. Therefore, it may be useful to note here that book authors have very limited or no influence on the decisions and actions of publishers.
Fifteen years after the publication of Braem’s first book dealing with the genus Paphiopedilum, Braem & Chiron presented a new monograph, and now, another 10 years later, we present again an updated edition, this time as an e-book with the help of Mr. Lars Pedersen of Moorland eBooks. Sandy Öhlund joined us as a co-author, adding valuable contributions regarding Paphiopedilum growing and judging, and Eric Sauer generously agreed to write a comprehensive treatment of Paphiopedilum culture.
It is simply impossible to write a book that satisfies everyone. The book has to be as complete as possible within the limits of technical and commercial feasibility. There are some chapters that may not necessarily interest the “general public” but that are of primary importance to the botanist: for example, the chapters referring to the species concept, the problems of taxonomy, systematics, and nomenclature, the chapters regarding the Orchidales and the slipper orchids in general, as well as the more scientific chapters regarding the genus Paphiopedilum. We have attempted to give the book a layout which allows anyone to go directly to any specific part.
We sincerely hope that this monograph will add to the understanding and enjoyment of the genus Paphiopedilum.
Prof. Dr. Guido J. Braem - Lahnau, Germany, 2013
Dr. Guy Chiron - Voreppe, France, 2013
Sandra Öhlund - Rolling Prairie, Indiana, USA, 2013
GENERAL SECTION
INTRODUCTION TO THE TAXONOMY AND NOMENCLATURE OF THE GENUS PAPHIOPEDILUM
Taxonomy is part of biology. It is the study of classification, including its rules, theories, principles, and procedures. Plant taxonomists have been trained to identify plants and to classify them into a hierarchical system. Furthermore, if they come to the conclusion that the organism under evaluation is new to science, they have learned to prepare and publish a scientifically correct description with a name for the new plant. A classified organism or a group of classified organisms thus becomes a taxonomic unit and is referred to as a taxon (plural taxa). A taxon therefore can be a family, a genus, a species, etc. To do such classification, the taxonomist uses a number of techniques, covering, in effect, nearly the entire scope of the biological sciences (anatomy, morphology, biochemistry, cytogenetics, electron microscopy, etc.). Thus, in order to be a good taxonomist, one must master these fields and de facto be a good general biologist. But the work of a modern taxonomist does not stop here. It continues into the study of the interrelationships among the various taxonomic entities (taxa). This newer form of taxonomy, which is not limited to classification, is usually called systematics. The two terms, however, are often used interchangeably.
Biological research is neither restricted to a single country nor to a single language area. Thus, a taxonomist must have an understanding of various languages. It is not acceptable to do taxonomy based only on material published in one or two languages or in any given country. Of course, one cannot expect all taxonomists to master all languages. That would be nice, but hardly achievable within the limited scope of the human lifetime. However, not mastering a language is by no means an excuse for neglecting work published in that language, and no taxonomist, or anyone for that matter, ought to be ashamed to solicit the help of a translator to help him/her comprehend the contents of a foreign-language article. Misinterpreting materials often leads to misunderstandings and erroneous conclusions.
We have talked about books and periodicals, and this may already indicate the importance of literature sources for the taxonomist and taxonomy. One journal and a few general books that discuss orchids in a more or less superficial way are simply not enough, but most orchid hobbyists, in all countries, are limited to just that.
So, what are the requirements for good taxonomy? The answer is simple: a well-trained botanist who has studied systematics, who speaks several languages, and who has access to the pertinent literature.
Unfortunately, taxonomy is the most underrated and misunderstood branch of the biological sciences, even though it is probably the most important. Of course one may argue that it is possible to study any aspect of biology without knowing the identity of the organism under examination, but just try to publish your results ... well, one can always refer to the object of study as “the thing” ...
The following anecdote is offered to illustrate this situation.
An American university professor once found a hand-written note pinned to the door of his office. It read:
10:38 AM 5/22
In Bio. faculty Library 2nd floor until 2 PM today
(tall guy in green teeshirt)
Dear Sir,
I need a one hour lesson in keying out plants of the Cascades.
I have a job in which I am looking for 25 “endangered plants” in genie (sic) of Eriogonum, Delphinium, Hackelia, Silene, Astragalus, etc
I will pay $ 50.00 for this lesson ... I must leave for this job by tomorrow afternoon
(...) 321-6505
We do not know whether the professor ever contacted the “guy in the green teeshirt”, what was said if they did meet, or how far the student’s career went in the biological sciences.
PROBLEMS IN BOTANICAL TAXONOMY
The great majority of those who read this book will not be familiar with taxonomic questions in general. So we are presenting a short discussion of some of the taxonomic principles insofar as they are pertinent to our treatment of the genus Paphiopedilum. Those who are interested in obtaining in-depth knowledge of all aspects of taxonomy are advised to obtain a textbook dealing with that subject.
The Orchidaceae are generally classified within the monocotyledons (generally referred to as “monocots”). Monocots are plants which, upon germination, produce a single seed leaf (cotyledon) that provides nourishment for the germinating seedling. This classification is not perfect as only some orchids produce cotyledon-like structures. Hence, the classification of the Orchidaceae as a “monocot” family finds its reason in practicality and convenience only. The alternative would be to create a third “division” since the currently accepted categories are monocots and dicots (plants generating two seed leaves upon germination).
THE SPECIES CONCEPT
In a letter to the American botanist Asa Gray, dated 20 July 1857, the great Charles Darwin wrote that he had come to the conclusion that species are but strongly defined varieties. The species concept was by no means solved by that statement, and in the plant kingdom the issue has been a special problem for many years and still remains unresolved. Numerous definitions of “a plant species” have been phrased. All of them are extremely arbitrary and include terms that are open to interpretation. A few are more or less usable, but even those contain a number of criteria which require further clarification. The reason for this problem is that there are no generally accepted rules for the delimitation of a species or any other taxon. Hence, it is quite possible that different authors accept different criteria to differentiate among taxa at any level.
The only generally applicable definition of a plant species is: “A plant species is that which a competent botanist considers it to be.” (A.J. Richards, personal communication, 1982)
In Paphiopedilum, most species, such as Paphiopedilum delenatii, P. sanderianum, P. spicerianum, P. druryi, P. sangii, P. canhii, and P. henryanum, etc., present no problem in respect to taxonomy. The overall structure and morphology of their flowers is so unique that they are easy to identify even though they show some variation.
Others, however, do present serious problems, and they cannot be readily identified as valid species. For example, there are problems with Paphiopedilum gardineri and P. roebbelenii, as well as with several taxa in the subgenus Cochlopetalum. In these cases, the interpretation of each and every one of these taxa is very subjective and depends on where one draws the line regarding how much variation is reasonable within a naturally occurring entity. For this reason, we have combined some debatable taxa into complexes, and we leave it to our readers to decide for themselves what interpretation of the term “species” best suits their understanding and/or meets their needs.
For the same reason, we often use the word taxon (singular) or taxa (plural) instead of species. Referring to the taxon Paphiopedilum roebbelenii, for example, leaves room to interpret it as a separate, autonomous species, or as a variant (at whatever infra-specific level) of Paphiopedilum philippinense.
Most of the problems we encounter, however, are man-made. Some, but by no means all, are discussed in the following sections.
TRAINING
Most orchid taxonomy is done by people who have little if any adequate training in botany. Whereas no one would dream of letting a baker or a plumber take out his/her appendix, we find that many in the orchid community feel called upon to do orchid taxonomy. Some of these people, in fact, do produce fairly decent to good material. Many, however, haven’t the faintest idea what they are talking and/or writing about. And that is not a surprise. After all, they do not have the training or the plant resources or the library or the ability to use them. These people may be very good in their learned professions, but that does not mean they are competent in the field of taxonomy.
MISIDENTIFICATIONS
Very often, commercial growers import plants without flowers and rely on the information they receive from their supplier or collector. Thus, whatever plant is supplied as “Paphiopedilum lowii”, for example, is labelled, treated, and sold as “Paphiopedilum lowii“. If, upon flowering, this proves to be wrong, the name is usually, but not always, corrected in the nursery. The clients, however, who have already obtained one of these plants, often do not recognise the mistake. They are, of course, not notified by the supplier, and end up with a misidentified plant in their collection. Thus if one calls attention to a misnamed plant in a hobbyist greenhouse, one is often told, “Well, that is what so-and-so says it is.” or “Well, that is what the tag says.” This especially applies when a plant has received a horticultural award. Good examples of this are Paphiopedilum stonei var. latifolium ‘Ruth Kennedy’, Phragmipedium schlimii ‘Wilcox’ and Phragmipedium schlimii ‘Birchwood’. These were misidentified decades ago and awarded under their false names by various orchid societies. The fact that “Paphiopedilum stonei” var. latifolium has very recently been described as an autonomous species has not changed and will not change matters much. More recently we saw the case of a very renowned California orchid nursery selling thousands of seedlings of the primary hybrid Paphiopedilum Prince Edward of York (P. sanderianum x P. rothschildianum) as pure P. sanderianum. The debacle is known among insiders, but one can be certain that many of these seedlings, now mature plants, are part of collections as P. sanderianum and that some of those might even be used in hybridisation. As of March 2013 at least one German commercial nursery still sells these plants as P. sanderianum, and on the website of a horticulturist for Pinecrest Gardens in Florida, a plant of P. Prince Edward of York is being depicted and praised as P. sanderianum.
BAD TYPIFICATION
Botanists assign a so-called type specimen to each taxon (family, genus, species, subspecies, variety, form, etc.). The designation of a type specimen is a prerequisite for a valid description according to the International Code of Nomenclature for Algae, Fungi, and Plants (formerly: International Rules of Botanical Nomenclature), generally referred to as the “Code”. Typification is one of the six principles of the Code. A type specimen is generally a plant, or a part thereof, which was deposited in an herbarium. In principle, the type should allow for a positive identification of the taxon that has been described. It can be a dried plant, or a dried flower. However, dried specimens are very often useless as they are many times damaged beyond proper recognition during the drying and mounting process. Typically collections of dried plants glued to paper are prime habitats for lower organisms that have cellulose on their favourite menu. This material, therefore, is only useful for taxonomy if the specimens are kept in proper cases and proper facilities. Unfortunately, this is something most botanical institutes cannot afford to do. Herbarium collections need to be cared for, but again, most institutes are understaffed and under-budgeted (the former, of course, generally being a result of the latter). Alcohol specimens are an excellent alternative to the “classical” dried specimens. They have many advantages, but they too have their problems. Whereas, for example, one will have no trouble finding a suitable receptacle to harbour a flower of Masdevallia, Oncidium, or most Paphiopedilum species, one will undoubtedly be challenged if one is to look for a receptacle to accommodate an entire plant of, say, Paphiopedilum sanderianum. And just imagine trying to conserve a plant of Grammatophyllum with its 2.5 m long pseudobulbs and 3 m long inflorescence in alcohol. But having overcome this “minor” problem, we will soon realise that alcohol specimens require special care, also. As there is leakage, the volatile alcohol diminishes constantly and needs to be replaced. The bottles must be stored properly. Another important drawback is that the alcohol mixture generally used to preserve the specimen contains formaldehyde, a carcinogenic component, and is highly flammable. Therefore, it must be handled with great care and in compliance with insurance and fire department rules.
The great advantage of the storage of specimens in alcohol is that they retain their form. If you don’t think this to be important just try to restore the staminode of a slipper orchid from a dried specimen! However, alcohol specimens decolourise quite rapidly. For that reason, they should always be paired with a colour photograph of the flower. Unfortunately, this is not required by the Code and almost never done.
Sometimes these types are very useful, quite often they merely add to the confusion surrounding a given taxon, and in some cases they may even be the primary cause for such confusion. As we have already stated, type specimens can be damaged beyond certain recognition. They may be lost or destroyed. During one single bombing raid on Berlin in World War II, for example, nearly the entire Schlechter Herbarium was lost. Another problem case is the Reichenbach Herbarium in the Vienna Museum of Natural History. H.G. Reichenbach’s herbarium and library were left to the Natural History Museum in the city of Vienna on the condition that “the preserved Orchids and drawings of Orchids” were not to be consulted during the first 25 years after his death. Thus, Reichenbach’s types were inaccessible between 1889 and 1914. Of course, Reichenbach’s decision was met with the most various reactions from his colleagues, ranging from disbelief and anger to dismay. Joseph Dalton Hooker claimed that Reichenbach had promised the herbarium to Kew and expressed great disappointment (although he had always kept Reichenbach from publishing in Curtis’s Botanical Magazine and other Kew-controlled publications). C.A. Backer (1936) may have been close to the truth when he wrote: “And so not mindful of the lesson in Galat. 5, 26 ‘Let us not seek vain glory’, in order to remain the mourned specialist himself, he made work difficult for others for a quarter of a century after his death.”
H.G. Reichenbach (generally referred to as Reichenbach fil.) is infamous among botanists for his short (to extremely short) and vague descriptions that in many instances simply may be regarded as useless, especially when the specimens upon which the descriptions are based are not available. Add to the 25 years of “unavailability by testament” the years of World War I and the result is that the Reichenbach material (including many types) was not accessible for at least one entire generation of botanists. Another problem is that sometimes reference is made to types that have never reached the herbarium, for whatever reasons.
If no type material can be located the rules of taxonomy call for the designation of new plant material which is then called a neotype. Too many neotypes, however, are subject to interpretation as they can be designated by any person who discovers that a type specimen is missing. A neotype, therefore, often mirrors that person’s interpretation of the original description. That original description itself, as we have seen above, can be very incomplete or dubious. Thus, it is often difficult, if not impossible, to be absolutely certain whether the plant being studied really corresponds to the plant which was available to the author of the original description.
Some botanists, especially those working in herbaria, consider these specimens to be the ultimate and only valid source for determining the identity of a plant. They often overlook the fact that in many cases there is no proof whatsoever that the designated “type” is, in fact, identical with the described plant. A good example of this is Paphiopedilum elliottianum, for which the supposed type specimen was produced several years after the original description!
The conclusion must be that herbarium materials, while sometimes very valuable and informative, should not be overrated and should never be relied upon as the only source of information about a plant.
SYNONYMS
Many taxa have several Latin names. There may be various reasons for this. The problem generally occurs when minor or geographical variants are described as species. This is nothing new and cases are known from such famous botanists as Lindley, Blume, Reichenbach, and Schlechter. Commercial growers obtain more revenue from a plant when it is called a “new” species than from that same plant designated as a variety. Thus, plants showing the slightest variation in leaf tessellation and/or flower colour often are given a different name and published as a separate species. Plants also obtain a new name when they are transferred to a new genus. Most Paphiopedilum species, for example, were originally published as Cypripedium species, and at one stage, nearly all of them were transferred to the genus Cordula by Rolfe. We will re-encounter these names in the descriptive part either as basionyms or synonyms. Sometimes, it is discovered that a designation is invalid. In those cases, a new name must be assigned according to the rules. In other cases, the same plant may be described independently by different authors. One example of this is the confusion and ongoing discussion regarding the names Paphiopedilum markianum and P. tigrinum. A second example may be found in the three independent publications of the same plant as Paphiopedilum vietnamense, P. mirabile, and P. hilmari. But this phenomenon of multiple descriptions cannot be eliminated. In a field where the nomenclature of a taxonomic group is based upon priority of publication, one can hardly blame anyone for trying to publish his or her concept first. Surely, very few authors inform their colleagues about what they are working on, and/or what they are about to publish, well knowing that if they did, others would try to publish more rapidly. One would be very naive if one were to believe the field of orchid taxonomy to be without professional jealousy which in recent years has also shown nationalistic features. Indeed, some people do believe that plants native to any given country should be described only by authors of that country. In respect to Paphiopedilum, this syndrome is well-known from China.
Our knowledge of the genus Paphiopedilum is constantly increasing. New species will be discovered and described. Literature searches and review of old materials may reveal that well-known names are invalid or have been used incorrectly; familiar names may disappear and new ones may need to be added to the inventory of the genus (as in the case of Paphiopedilum callosum, now correctly designated as P. crossii). Taxonomy is not a static part of science, and orchid taxonomy is by no means an exception.
ORCHIDACEAE AKA ORCHIDS
GENERAL CONSIDERATIONS
Since the publication of Carolus Linnaeus’ Species Plantarum in 1753, the taxonomy of all living matter has been classified mainly according to sexual characteristics. In flowering plants, these characteristics are found in the flowers, or in other words, in those structures that have been adapted for sexual reproduction that causes variation, thus producing the basis for evolution.
Until about forty years ago, students of biology had an easy task of dividing the living matter on our planet as either plants or animals. Those organisms having a cell wall were classified as plants and those without a cell wall were considered to be animals.
Since then, the world around us has been reclassified, and scientists now generally divide living things into six “kingdoms”: Bacteria; Archaea; Protista; Fungi (covering moulds and mushrooms); Animalia (multicellular organisms with cells that have an organised nucleus but no cell wall); and Plantae (multicellular organisms with cells that have an organised nucleus and a cell wall).
The plant “kingdom” is divided into many different parts, of which the flowering plants, scientifically referred to as Angiosperms, are but one. Within this large group (about 400 families with a total of about 250,000 species), the Order Orchidales is of special interest to us and will be the focus of more extensive discussion.
THE ORCHIDS
The orchids, being one of the largest and most diverse groups of the plant “kingdom”, comprise between eight and ten percent of all known flowering plants. This represents between 25,000 and 35,000 taxa. A total of 59,695 entries of Orchidaceae names were included in the Index Kewensis2, up to June 1996. As of 29 September 2012, the number of entries had increased to 77,915. This number, however, does not give an accurate count of the number of species because it includes subspecies, varieties, forms, as well as synonyms. Orchids are an actively evolving group with extremely specialised flowers that are adapted for attracting, deceiving, and manipulating insects to achieve cross-pollination.
The following characteristics are common to all plants in the orchid family. Some of these characteristics, however, are shared by other groups of plants.
(1) The inflorescence is either terminal or lateral, meaning that it is generated from the apex or from the side of a plant growth.
(2) The ovaries are inferior, meaning that the ovary is below the stamens, sepals and petals.
(3) The flowers are bilaterally symmetrical, meaning that they can only be split into similar halves along one given plane.
(4) There are six tepals in two whorls.
(5) The three tepals of the outer whorl are usually referred to as sepals. The three tepals of the inner whorl are called petals.
(6) The median tepal of the inner whorl is referred to as the labellum or “lip”.
(7) The number of stamens is always reduced.
(8) The median stamen of the outer whorl and the lateral ones of the inner whorl are developed as either fertile stamens or as more or less distinct staminodes.
(9) Stamens and style are fused into a gynostemium which forms a “column” except in a few genera.
(10) The gynostemium of the orchids is a unique feature within the monocots, but analogous structures are found in two families of the dicots, namely the Stylidiaceae (Trigger Plants) and the Asclepiadaceae (Milkweeds).
(11) The anthers are joined to the filament at its base (basifixed) or for some distance along its dorsal edge (dorsifixed).
(12) The pollen grains are single in the Apostasioideae and Cypripedioideae, but cohere in tetrads in the majority of all other orchids.
(13) The ovary may be unilocular, with a single chamber in which the pollen grains develop, or trilocular, with three such chambers. There is wide disagreement about the interpretation of what can be deduced from a cross-section of an orchid ovary (Brown, 1833; Vermeulen, 1966).
(14) The style is more or less apically inflexed and terminated by a tri-lobed stigma with a sticky surface.
(15) The development of the embryonic sac is triggered by pollination. Sometimes, the actual fertilisation does not occur until five or six months after pollination (Wirth & Withner, 1959).
(16) The embryo is always immature in the ripe seed.
(17) The seeds are very minute and numerous, but vary considerably in shape and size.
(18) Upon germination the embryo forms a tubercle (protocorm) which is covered with hair-like structures on most of its basal part. Eventually several leaves develop from the upper end of the protocorm.
(19) Under natural conditions, most orchids will germinate only when a symbiosis with a fungus has been established. The adult green plants are usually viable without the presence of the fungus. The relationship of orchids to fungi is still under study and interpretations vary considerably.
The “construction” of the flowers, the numerous small seeds, and the close relationship with the fungi are characteristics that are shared between the orchids and the Burmanniales, a group of poorly understood, mainly tropical and subtropical monocots of no horticultural merit. For that reason, both groups are sometimes classified into one complex.
Scientifically verifiable facts about the evolution of the orchids are very rare. Unfortunately, the fossil record is very poor in respect to plants. The only fossil hitherto found that can be unambiguously linked to an orchid is the discovery of an orchid pollinarium attached to the mesothorax of an extinct stingless bee (Proplebeia dominicana), preserved in Miocene amber from the Dominican Republic (Ramírez et al., 2007). The fossil was dated to be between 15 and 20 million years old. It is assumed that orchids are probably derived from lily-like ancestors with six stamens. The orchid genus Neuwiedia may be a relic of an undetermined earlier stage. But this assumption should be regarded as an intelligent guess, and other interpretations are quite possible.
Whereas there is general agreement on the major groups of orchids in respect to their broad outline, there is much disagreement about their ranking and the relationships among them. Many modern treatments follow Garay (1960, 1972) in considering the orchids as consisting of one family, containing five or six subfamilies. However, as early as 1833, John Lindley, the “Father of Orchidology”, suggested recognising three families within the orchids. Lindley did not discuss this division in detail, but the text in his Nixus plantarum is quite self-explanatory. This part of Lindley´s work has been generally neglected. Vermeulen (1966) was the first to resurrect the idea, and Dahlgren, Clifford, & Yeo accepted this division for The Families of the Monocotyledons, published in 1985. The actual division is made among the Apostasiaceae, Cypripediaceae, and Orchidaceae. The differences can best be taken from the following key:
It should be noted that, no matter what approach one adopts - three families or a single family with a number of subfamilies - the result is exactly the same. The position of the various taxa is merely shifted by one level within the taxonomic hierarchy.
THE CYPRIPEDIAE
The Cypripediae or “slipper orchids” were among the first orchids mentioned at the generic level (Heister, 1748). In 1753, Linneaus, the founder of modern taxonomy, created, among others, the genus Cypripedium to accommodate Cypripedium calceolus, the only slipper orchid present in central Europe.
By the end of the 19th century, the genus Cypripedium L. included about one hundred species. Of these, approximately twenty-five percent were known from the temperate regions. The rest originated from the Asian and American tropics and sub-tropics.
In 1818, Rafinesque created the genus Criosanthes with the type Criosanthes arietinum, a North American slipper orchid. Twenty years later, in volume four of his Flora Telluriana, he founded three new genera for Asian slipper orchids: Stimegas with the type species Stimegas venustum, Cordula with the type species Cordula insignis, and Menephora with Menephora bicolor as its type. Rafinesque’s intentions were rather clear in describing these taxa. He wrote (loc. cit. sub No. 931): “I propose the following subgenera or rather genera ...” and, although his descriptions are very short, his genera Stimegas and Cordula were clearly typified. We do not know, however, what Rafinesque meant with his genus Menephora. His description is not adequate to clarify the identity of the plant designated as Menephora bicolor. The description states: “ ... Borneo and Java ... flowers dull purple, but upper petal white, broader ovate.” It has been suggested that Menephora bicolor is synonymous with Paphiopedilum purpuratum but this latter species does not occur in either Borneo or Java. One explanation is that Rafinesque referred to a plant of the Paphiopedilum javanicum complex. Another possibility is that Rafinesque erred about the natural distribution of the plant now generally known as P. purpuratum.
Until about forty years ago, the work of Rafinesque was not taken into consideration by most authors. Some just ignored it, others (the majority) didn’t know of his writings. Indeed, only a few copies of his works have survived in specialised libraries. Upon Rafinesque’s death in 1840, the remaining stock of his books was sold as “scrap paper”.
In 1848, Lindley founded the genus Uropedium based on materials from the Linden expedition to the American tropics, and in 1854, another genus, Selenipedium, was described by H. G. Reichenbach (Reichenbach fil.) to accommodate two other slipper orchids from that continent. Uropedium as well as Selenipedium suffered a fate similar to that of Rafinesque’s work: they were, for the most part, generally ignored.
Pfitzer, a German botanist working in Heidelberg, separated all the tropical species from Cypripedium and combined them in a new autonomous genus Paphiopedilum (1886). He included a section Phragmopedilum for the species with conduplicate leaves from the American tropics, described as Selenipedium section Acaulia by Reichenbach fil.
In 1896, Rolfe elevated Pfitzer’s section Phragmopedilum to the genus Phragmipedium in his review entitled The Cypripedium Group. Rolfe differentiated among the genera Selenipedium Reichenbach fil., Phragmipedium Rolfe, Cypripedium L., and Paphiopedilum Pfitzer. In 1901, Rolfe changed the name of the genus Phragmipedium to “Phragmopedilum” as he considered the latter a more linguistically correct spelling. However, the rules of nomenclature do not permit such corrections, no matter whether they are linguistically indicated or not.
In the meantime, a rather extensive study by Johannes Gottfried Hallier (Hallier fil.) was published in the Annales du Jardin Botanique de Buitenzorg. In that study, Hallier (1897) discussed a new species, Paphiopedilum amabile, and presented an “overview” of the entire genus.
Pfitzer (1903) again reviewed all slipper orchids for Engler’s series Das Pflanzenreich. Pfitzer based this revision on Hallier’s work of 1897 and recognised Selenipedium Reichenbach fil., Phragmipedium Rolfe as Phragmopedilum (Pfitzer) Rolfe emendates (= corrected) , and Paphiopedilum Pfitzer as separate genera. Pfitzer must have been aware of at least part of Rafinesque’s work as he reduced the Rafinesque genus Criosanthes to a section of the genus Cypripedium L. He did not, however, mention Rafinesque’s genera Stimegas, Cordula, and Menephora.
In 1959, the International Botanical Committee conserved Paphiopedilum against the earlier published Cordula and Stimegas (see below).
In 1971, at the beginning of the publication of the 3rd edition of Rudolf Schlechter’s Die Orchideen, Brieger made a revision of the slipper orchids. Brieger divided the subfamily into four tribes, each with one monogeneric subtribe. On the generic level, Brieger recognised Selenipedium Reichenbach fil, Phragmipedium Rolfe, Paphiopedilum Pfitzer and Cypripedium L.
In 1975, Phragmipedium was conserved over the earlier published name Uropedium at the botanical congress in St. Petersburg (then Leningrad), following a proposal by Dressler & Williams.
The relationship among the slipper orchids was then the subject of an extensive study by John T. Atwood (1984). One of the conclusions he reached was that the plant generally known as Cypripedium arietinum differs quite distinctly from all other species within the genus Cypripedium, and also shares two anatomical features with the genus Selenipedium. He therefore “revived” the genus Criosanthes.
Albert & Chase (1992) founded the genus Mexipedium to accommodate a plant originally described as Phragmipedium xerophyticum.
The genera Criosanthes Rafinesque and Mexipedium Albert & Chase are not generally accepted. Up to this time, most authors have considered Criosanthes part of Cypripedium. Albert (1994) used cladistic analysis of morphological and molecular data to reconfirm that Cypripedium, Paphiopedilum, and Phragmipedium, including Mexipedium and Selenipedium, are derived from a common ancestor. Albert & Pettersson (1994) suggested placing Mexipedium and Phragmipedium back into the genus Paphiopedilum. These suggestions are based on data obtained using methods of molecular biology which are enormously overrated and should not be used out of the context of the entire suite of taxonomic methods, especially those now classified as alpha-taxonomy.
As one would expect, plants sharing common ancestors should have some characteristics and a number of genetic sequences in common. All that is needed, however, is to put a plant of each of the different genera of the slipper orchids together on a table or greenhouse bench, to make obvious what immense differences there are among them.