SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 61 

A Phylogenetic Analysis 

of the-Orchidaceae 

Pamela Burns-Balogh 

and V.A. Funk 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1986 
ABSTRACT 

Burns-Balogh, Pamela, and V.A. Funk. A Phylogenetic Analysis of the 

Orchidaceae. Smithsonian Contributions to Botany, number 6 1, 79 pages, 94 

figures, 4 plates, 1986.-The Orchidaceae is the largest flowering plant 

family, with approximately 25,000 species. Sixty-eight apomorphies grouped 

into forty-two transformation series were used to construct a cladogram for 

the twenty-six tribes of the family. A detailed discussion of the characters is 

followed by an in-depth analysis of the cladogram. The cladogram was used 

to develop a classification and a natural key to the tribes. Seven subfamilies 

(Neuwiedioideae, Apostasioideae, Cypripedioideae, Spiranthoideae, Neottioideae, Orchidoideae, and Epidendroideae) are divided into 20 tribes, the 

majority of which are defined by synapomorphies. Of the intrasubfamily 

classifications, that of the Epidendroideae is the most tenuous. The phylogeny 

and classification presented here are hypotheses of relationship and are 

therefore subject to change as more information becomes available. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is 

recorded in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: Leaf 

clearing from the katsura tree Cerecidiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Burns-Balogh, Pamela. 

A phylogenetic analysis of the Orchidaceae. 

(Smithsonian contributions to botany ; no. 61) 

Bibliography: p. 

Supt. of Docs. no.: SI 1.29: 61 

1. Orchids-Classification. 2. Cladistic analysis. 3. Plants-Phylogeny. I. Funk, V. A. 

(Vicki A.), 

1947- . 11. Title. 111. Series. 

QKl.S2747 no. 61 581s [584’.15’012] 85-600315 [QK495.064] 
Contents 

Page 

Introduction 1 ............................................ Acknowledgments .................................... 1 

History of Orchid Classification ............................ Objectives of Systematics ................................. 

Principles of Phylogenetic Systematics ..................... Traditional Orchid Systematics .......................... Phenetics and Orchid Classification ....................... 

Cladistic Analysis of the Orchidaceae ......................... 

Synopsis of Burns-Balogh and Funk Classification ............ 

4 

5 

5 

6 

8 

9 

Character Selection ..................................... 9 

14 

Discussion of Characters and Their Aggregation into Transformation Series ........................................ 15 

Characters of the Pistil ................................. 15 

Characters of the Stamen ............................... 19 

Characters of the Pollinarium ........................... 30 

Characters Not Used in this Classification .................. 40 

Cladogram Construction ................................ 40 

Discussion of Characters as Displayed on the Cladogram ......... 41 

Discussion of the Cladogram and Resulting Classification ........ 45 

Keys to the Subfamilies and Tribes of the Orchidaceae ........... 50 

Classification of the Orchidaceae ............................ 53 

Conclusion ............................................. 60 

Appendix-List of Specimens Examined ...................... 62 

Literature Cited ......................................... 70 

Platesl-4 .............................................. 72 

... 

111 

A Phylogenetic Analysis 

of the Orchidaceae 

Pamela Burns-Balogh 

and V.A. Funk 

Introduction 

Traditionally, the orchids have been separated 

into two major groups (formal or informal) based 

on the number of anthers: monandrous and nonmonandrous (Pfitzer, 1888- 1889; Schlechter, 

1926). The monandrous orchids (one functional 

stamen) comprise the largest group, with approximately 25,000 species, and consist of the 

subfamilies Spiranthoideae, Orchidoideae, Epidendroideae and Vandoideae (for this section of 

the paper all categories are those of Dressler 

1981; see Table 1). The second group contains 

orchids that have more than one functional stamen. This group is much smaller, with about 200 

species in two subfamilies, the Apostasioideae 

and Cypripedioideae. Because of the stamen 

number and partial column development, the 

orchids with more than one anther (misleadingly 

referred to as diandrous) are considered to be 

more primitive than the monandrous orchids 

(e.g., Dressler, 198 1). The categories within the 

subfamilies (tribes and subtribes) are based on a 

variety of characters including vegetative and 

floral features and habit type. As is true with a 

number of the flowering plant families, many of 

the groups (subfamilies, tribes and subtribes) proPamela Burns-Balogh and V.A. Funk, Department of Botany, 

National Museum of 'Vatural History, Smithsonian Institution, 

Washington, D.C. 20560. 

posed in the various classifications of the orchids 

are poorly defined and subject to constant rearrangements that appear, at least in some cases, 

to be merely a matter of intuition. 

We have identified 68 apomorphies organized 

into 42 transformation series. The apomorphies 

were used to construct a cladogram (Figure 1) 

from which a classification was developed. The 

classification presented here recognizes groups 

that the cladogram indicates to be natural or 

monophyletic (sensu Hennig, 1966). The characters used to develop the cladogram were those 

we found to be consistent within the monophyletic groups they are inferred to delimit (in most 

instances this was the tribal level). The information available on the column structure for members of the Epidendroideae is incomplete, and 

consequently the resulting classification for that 

subfamily is somewhat tentative. The classification presented in this paper is compared in Table 

1 with those of Dressler (1981), Garay (1972), 

Schlechter (1 970- 1984), Vermeulen (1 966), 

Dressler and Dodson (1 960), and Rasmussen 

( 1 98 3). 

Robinson, R.L. Dressler, F.N. Rasmussen, C.J. 

Humphries, E.W. Greenwood, R. Read, W. 

Barthlott and five anonymous reviewers for their 

ACKNOWLEDGMENTS.-We are grateful to H. 

1 
2 

Taxonomist 

Balogh and Funk (1 985) 

Dressler and Dodson (1 960) 

Vermuelen (1 966) 

Schlechter (1970-1984) 

Garay (1 972) 

Dressler (1 98 1) 

Rasmussen (1 983) 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 1 .-Classification systems of Orchidaceae 

Apostasioideae C ypripedioideae Spiranthoideae 

Apostasioideae Cypripedioideae Spiran thoideae 

Neuwiedioideae Cranichideae 

Prasophylleae 

Diurideae 

Apostasiaceae 

Apostasiaceae 

Apostasioideae 

Apostasioideae 

Apostasiaceae 

Apostasieae 

Cypripedieae 

C ypripediaceae 

Cypripedioideae 

Cypripedieae 

Paphiopedileae 

Phragmipedieae 

Selenipedieae 

C ypripedioideae 

Cypripedioideae 

Cypripediaceae 

Spiranthoideae 

Erythrodeae 

Cranichideae 
NUMBER 61 

TABLE 1.-Continued. 

3 

Neottioideae Orchidoideae Epidendroideae Vandoideae 

Neottioideae Orchidoideae Epidendroideae - 

Neottieae Diseae Arethuseae S.S. 

Thelymitreae Satyrieae Vanillieae 

Geoblasteae Orchideae Gastrodieae 

Pterostylideae Triphoreae 

Epidendreae 

Pleurothallis Group 

Dendrobieae 

Malaxideae 

Maxillarieae 

Vandeae 

Coelogyneae 

- 

Epidendroideae 

Contribe Neottieae 

Neottioideae 

Tropideae 

Neottieae 

Spirantheae 

Diurideae 

Neottioideae 

Epipogieae 

Cranichideae 

Neottieae 

Orchidoideae 

Neottieae 

Orchideae 

Epidendreae 

Orchidoideae 

Orchidoideae 

Satyrieae 

Orchideae 

Diseae 

Orchidoideae 

Orchideae 

Diseae 

Disperideae 

Orc hidoideae 

Diseae 

Neottieae 

Diurideae 

Orchideae 

Orc hidoideae 

Diurideae 

Orchideae 

Epidendroideae 

Contribe Epidendranthae 

Epidendroideae 

Podochileae 

Arethuseae, 

Epidendreae 

Vandoideae 

Maxillarieae 

Oncidieae 

Cataseteae 

Vandeae 

Epidendroideae 

Epidendreae 

Vandeae 

Vandoideae 

Cymbidieae 

Vandeae 

Pol ystachyeae 

Maxillaireae 

Epidendroideae 

Epipogieae 

Vanillieae 

Gastrodieae 

Arethuseae 

Coelogyneae 

Malaxideae 

Cryptarrheneae 

Epidendreae 

Cal ypsoeae 

Neottioideae Epidendroideae Vandoideae 

Epipactieae Arethuseae Pol ystach yeae 

Neottieae Vanilleae Cymbidieae 

(= Spiranthoi- Gastrodieae Maxillarieae 

deae and Epipogieae Vandeae 

Listerinae) Coelogyneae 

Malaxideae 

Epidendreae 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

advice and criticism. We thank H. Robinson for 

his encouragement. We thank E. Greenwood, 

J.B. Comber, and M. Clements for the use of 

some of their color photos and the following 

for sending flowers, pollinaria, photos or drawings: P. Bernhardt, M. Clements, J.B. Comber, 

R.L. Dressler, E. Greenwood, E. Hagsater, H. 

Kurzweil, J. McQueen, and F.N. Rasmussen. 

Thanks go to B. Kahn for helping to assemble 

the manuscript. We also thank the National Museum of Natural History, Smithsonian Institution, Washington, D.C., and the University of 

Maryland, College Park, Maryland, for supplying 

SEM time and assistance. Fresh flowers were 

obtained from the US. National Orchid Collection and Marie Selby Botanical Garden. 

History of Orchid Classification 

John Lindley has aptly been named the father 

of orchidology. The work that eventually became 

his book, The Genera and Species of Orchidaceous 

Plants (1830-1840), is the basis on which all 

modern systems of orchid classification are 

founded. He divided the approximately 2000 

species of orchids then known into seven tribes 

on the basis of anther number and type of pollinarium. A variation of Lindley’s system appeared 

during the latter part of the 1800s in Bentham 

(1881). In 1888 and 1889, Pfitzer used vegetative characters as well as column characters to 

separate the orchids into groups. His major divisions were based on the following: number of 

anthers, stem and leaf characters, and the consistency of the pollinaria. For many orchidologists his system was not satisfactory because many 

genera considered to be closely related were put 

in separate alliances. Later, Rudolf Schlechter 

(1 926) developed a system of classification modified from Pfitzer. Schlechter’s system is still used 

in some major herbaria. His tribes were based 

on anther and pollinarium characters while the 

subtribes were based on vegetative and floral 

characters. 

In 1937, Mansfield treated only the monandrous orchids using the methods of both Bentham (1881) and Schlechter (1926), but his 

groups were basically those of Lindley, although 

many of the genera had been placed in different 

groups. Dressler and Dodson (1 960) segregated 

the orchids into tribes and subtribes based on 

their level of specialization, but the classification 

still contained the principal groups devised by 

Lindley. 

There are two major conflicts that separate 

modern orchid systematists. The first difference 

of opinion concerns the level of recognition of 

some of the major taxa within the orchids. One 

group of workers emphasizes the similarities that 

unite all orchids (partial fusion of the filaments 

and style, lack of endosperm, mostly resupinate 

flowers, two inner lateral and one outer median 

stamen positions) and thinks the Cypripedioideae 

and Apostasioideae (diandrous and triandrous 

orchids) should be included in the Orchidaceae 

(Dressler, 1981; Garay, 1972). A second group 

of systematists stresses the importance of the 

differences between the diandrous/triandrous 

and monandrous orchids and raises one or two 

of the subfamilies mentioned above to family 

level (Vermeulen, 1966; Schlechter, 1926, 

The second major disagreement concerns the 

placement, recognition and contents of the 

subfamily Neottioideae (Garay, 1972; Polychondreae of Schlechter, 191 1). In 1826, Lindley 

described the Neottieae (sic; = subfamily), which 

included the tribes Neottieae and Gastrodieae. 

However, in a later treatment (1 830-1840) 

he dropped the higher category and only recognized the tribe Neottieae, which he characterized by the distinctive dorsal anther and the 

powdery pollen. Since Lindley’s latter work the 

genera in the Neottieae have been shuffled 

among the Epidendroideae, Orchidoideae, Spiranthoideae, and Neottioideae. Dressler and 

Dodson (1960) recognized the Neottieae as a 

tribe of the Orchidoideae; Vermeulen (1 966) 

recognized them at the contribe level (groups of 

tribes) and placed them in the Epidendroideae; 

Schlechter, (1 970-1 984) and Garay (1 972) recognized Neottieae as a distinct subfamily-Neot1970-1984). 
NUMBER 61 5 

tioideae. Dressler (1 98 1) recognized the significance of anther position when he reorganized 

and segregated the tribes in Neottioideae. He 

removed Erythrodeae and Cranichideae, because 

that group of orchids has inconspicuous staminodia and an erect dorsal anther subequal to the 

rostellum. He placed these two tribes in a new 

subfamily, Spiranthoideae. He placed the Diurideae and Neottieae with the Orchideae and the 

Diseae in the subfamily Orchidoideae because of 

their terminal erect anther, root-stem tuberoids, 

lack of stomata1 subsidiary cells, and a shared 

similar habit. Controversy arose when Dressler 

assumed the Orchidoideae (in the traditional 

sense) were specialized diurids. The placement 

of the “neottioid” orchids has received the recent 

attention of Rasmussen (1 982). He suggests that 

the neottioids are the lowermost branches of the 

monandrous orchids, i.e., a paraphyletic group, 

phylogenetically comprising all the other monandrous orchids. In his paper on the Orchidales 

(1 983) Rasmussen does include the Diurideae 

with Orchidoideae s. s. because of the scattered 

occurrence of orchidoid characters such as basitony. He regards Diurideae as a remnant group 

of Orchidoideae. 

Objectives of Systematics 

The way we “do systematics” is strongly influenced by what we hope to achieve with our 

systematic studies. Most taxonomic studies have 

two basic goals: (1) delimiting and identifying 

taxa, and (2) expressing the inter-relationships 

among the taxa. The process of achieving these 

goals can be referred to as developing a classification-certainly a good classification attains 

both these goals. There are different ways of 

developing a classification and the means one 

uses dictates the type of information expressed. 

One can be either intuitive or methodological; 

the latter procedure can be further expressed as 

either cladistics or phenetics. 

For a number of years orchid systematists have 

been relying on intuition; practically, this resulted in a combination of character weighting 

and some type of Gestalt evaluation (overall similarity). The taxonomist studied a group of organisms, selected characters she/he believed to 

be conservative (i.e., important), and delimited 

the groups of species based on these characters. 

Disagreements arose when a different taxonomist thought different characters were “more 

important.” If relationships within and among 

the groups of organisms were expressed, it was 

usually done with little explanation as to why 

certain relationships were selected. There have 

been two recent attempts to develop an empirical 

method for determining classifications-cladistics and phenetics. 

PRINCIPLES OF PHYLOCENETIC SYSTEMATICS 

The basis of phylogenetic systematics (Hennig, 

1966) is the recognition and maintenance of the 

phylogenetic relationships among monophyletic 

groups. The reasoning process works as shown 

in the following example. Given any three taxa, 

which two are more closely related to one another than either is to the third? The question is 

answered by finding an evolutionarily unique 

character(s) (an apomorphy) in two of the taxa 

and not in the third. The two that share the 

unique character are then grouped together. An 

apomorphy is called synapomorphy if it is shared 

by two or more taxa, or autapomorphy if it is 

found in only one taxon. Phylogenetic systematists choose to emphasize the geneological relationship among groups of taxa. We believe it is 

the best method for developing a classification 

(Farris, 1983; Nelson and Platnick, 198 1). Implementation of the method can be described generally as following these “steps:” select the characters, group them into transformation series, 

determine which character(s) in each transformation series is (are) apomorphic, group the taxa 

based on shared apomorphies, and use the resulting cladogram to develop a hierarchial classification. 

Hennig used the term phylogenetic systematics 

for his philosophy. The term cladistics, which 

was introduced later, is often used in place of 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

phylogenetic systematics. However, as defined 

by Wiley (198 l), cladistics is the method by which 

the cladogram is constructed and phylogenetic 

systematics is the entire philosophy of the 

method. Inherent in phylogenetic systematics are 

homology, parsimony, and monophylesis. 

Homology applies to characters inherited from 

a common ancestor. The entire character selection process is actually a search for homology. 

We attempt to put homologous characters into 

the same transformation series. For instance, we 

treat the hamular stipe and tegular stipe in two 

different transformation series because we have 

evidence from the studies of Rasmussen (1982) 

that they are structurally different. Of course, 

characters that appear to be homologous at the 

beginning of the study may be shown by the 

cladogram to be more easily explained with a 

hypothesis of parallel or convergent evolution 

(homoplasy), appearing two or more times. Concerning the stipe characters mentioned above, 

both were found at least two times on the cladogram (Figure 1) indicating that neither all of the 

tegular stipes nor all of the hamular stipes were 

homologous. 

Parsimony (accepting the simplest explanation 

for the data) is applied on two levels. On the 

character level it means that within each transformation series we orient the characters with 

respect to one another in the simplest way possible. This orientation is based on ontogenetic and 

developmental information for the study group 

and the characters found in the most closely 

related groups (outgroups). Those characters 

present in both the group under study and in the 

outgroup are said to be the general condition 

(Platnick, 1979) and are referred to as plesiomorphic (symplesiomorphies are shared general 

characters). The other characters within the 

transformation series are all considered to be 

apomorphic. Transformation series that have 

more than one apomorphic character are more 

difficult because it is impossible to tell from outgroup comparison how the apomorphies are related to one another. Because of this difficulty 

we have chosen to treat such apomorphies as 

independently derived. One of the more difficult 

cladistic concepts to explain is that apomorphies 

are relative. For instance, if we use the lack of 

endosperm as a synapomorphy for all orchids, 

then within the family that character is plesiomorphic. Likewise, if the presence of two inner 

lateral staminodes is a synapomorphy for the 

monandrous orchids then that character cannot 

be used to circumscribe groups such as Epidendroideae and Orchidoideae that are contained 

within the larger group of monandrous orchids. 

For the cladograms, applying the parsimony 

principle does not mean that evolution is parsimonious, it means that we seek the diagram that 

requires the fewest number of assumptions of 

parallel evolution and reversals (for a detailed 

explanation see Farris, 1983). Parsimony is the 

only logical criterion for choosing a cladogram 

because it gives us the one that best fits the data. 

Monophyletic groups are the only ones that 

are recognized as natural or significant in cladistic classifications. Monophyletic groups are delimited by the presence of an apomorphy. Nonmonophyletic groups are not definable by unique 

characters and have not had a unique history in 

common. Certainly non-monophyletic groups do 

not contain all of the descendants of an ancestor 

and are, therefore, not natural groups. 

Additional explanations on the above topics 

accompany the various sections of this study and 

can be found in the following references: Eldredge and Cracraft (1 980), Nelson and Platnick 

(1 98 l), and Wiley (1 98 1). 

We will first examine intuitive and phenetic 

classifications of orchids before presenting our 

cladistic classification. Previous to this study only 

a few cladistic studies of orchid genera have been 

published: Brownlea (Linder, 198 la), Monadenia 

(Linder, 1981b), Hershelia (Linder, 1981c), and 

the Neottioid orchids (Rasmussen, 1982). 

TRADITIONAL ORCHID SYSTEMATICS 

Orchid taxonomy is fraught with the classic 

problems of intuitive or Gestalt systematics. The 
NUMBER 61 7 

problems, which fall into a number of somewhat 

overlapping categories, include: the use of inconsistent characters, grouping by plesiomorphies, 

circular reasoning, intuition (used when there is 

a lack of data), appeal to authority, character 

weighting, overall similarity (phenetics), and the 

use of groups of characters as characters (a few 

examples are given below and additional discussion can be found throughout this paper). 

In 1981, Dressler used three characters to 

delimit the subfamily Orchidoideae: presence of 

root-stem tuberoids, lack of stornatal subsidary 

cells, and an erect anther. While Dressler placed 

all taxa that have root-stem tuberoids into the 

Orchidoideae, he also included taxa that did not 

have them, and in fact, this character is usually 

consistent only at the generic level. For instance, 

Rasmussen (pers. comm., 1983) reports that not 

all Diurideae, which supposedly have root-stem 

tuberoids, actually have this type of structure. 

His preliminary studies indicate this structure is 

only a swollen root without the polystelar rootstem tuberoid anatomy. The root stem tuberoid 

is absent in the Neottieae and is inconsistent as a 

character in the Orchideae. 

The erect anther is the plesiomorphic character (see “Discussion of Characters as Displayed 

on the Cladogram,” p. 41) giving rise to three 

different anther types: upside-down, at right angles to the column axis, and operculate-incumbent. The presence or absence of subsidiary cells 

is a very inadequately studied character (Rasmussen, 1982) and so much missing data limits 

its usefulness. While each of the three apomorphic characters can be used to group taxa, 

the retention of the plesiomorphic character 

gives us no information on relationship. Dressler’s Orchidoideae contains taxa with different 

column morphologies. He evidently considers 

the three characters listed above, two of which 

lack conclusive data, as more important; this is 

character weighting. 

Garay (1972) gives very little explanation as to 

why he groups certain things together in his 

classification, he simply presents it. This is a 

classic example of “appeal to authority” because 

we are asked to accept the classification because 

of expertise in the field, not because of data 

presented. Intuition and overall similarity work 

in much the same manner. Workers will cite 

Gestalt as a reason for separating taxa, or will 

discuss how different the subtribes are, but will 

not be able to list the characters they used to 

differentiate them. 

Apostasia and Neuwiedia are usually grouped 

together because they are primitive, and certain 

characters, such as partial fusion of the androecium and gynoecium, are considered primitive 

because they are found in these two genera. The 

fact remains, however, that many workers do not 

regard them as closely related (Garay, 1972). 

Garay stated that “Apostasia and Neuwiedia are 

not even closely related to one another but are 

only relic survivors of ancient anagenetic lines.” 

This is a fine example of circular reasoning. Why 

are the genera primitive? Because they have 

primitive characters. Why are the characters 

primitive? Because they are found in primitive 

groups. Actually we found that the two genera 

were not particularly closely related and that like 

all other taxa, both have some apomorphic and 

well as plesiomorphic characters. Another example of circular reasoning is found in Garay 

(1 960). He regards the heavy sclerotic seed coat 

in Vanilla as a primitive character because Apostasia nuda and Selenipedium chica have a similar 

type of seed. A character is not primitive because 

it is found in a primitive group. Our results agree 

with those of Rasmussen (1 983); the sclerotic 

seed coat is more likely secondarily derived. 

Often taxonomists will use characters that are 

really groups of characters; a good example in 

orchids is habit. Orchids are epiphytic, terrestrial, autotrophic, or saprophytic; much discussion takes place about which of these habits is 

primitive (Benzing, 1981). The habit is not a 

character. What are characters are the features 

that enable the plant to have one of the three 

habits, such as the presence or absence of velamen in the roots, or small seeds (Robinson and 

Burns-Balogh, 1982). 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PHENETICS AND ORCHID CLASSIFICATION 

Others have noted the various problems with 

traditional classification and have attempted to 

be more empirical in their studies. The various 

phenetic approaches concentrate on reflecting 

the total similarity of the organisms in question. 

In practice this means that given any three taxa 

the two that appear more similar to one another 

are grouped together. Lavarack completed a dissertation exploring the relationships within the 

Neottioideae (1 97 1, unseen) and published two 

papers summarizing his work (1974, 1976). He 

concentrated on the Australian taxa but the analysis included members of all of the subfamilies. 

In his 1976 paper he seeks to “consider relationships as indicated by an examination of overall 

similarity.” This type of study involves using all 

possible characters and evaluating the relationships using a variety of computer programs that 

use different algorithms (for a detailed discussion 

of the advantages and disadvantages of these 

programs see Sneath and Sokal, 1973). Although 

Lavarack used a few column characters for his 

published studies, he relied mainly on vegetative 

characters. 

Phenetic studies eliminate some of the problems of intuitive classification discussed above, 

such as circular reasoning, intuition, appeal to 

authority, and groups of characters. Other problems are retained, however, though sometimes 

in a different manner. For example, phenetic 

studies of orchids deal only with individuals and 

can therefore use characters that are known to 

vary within the group as a whole. For instance, 

one character in Lavarack’s study is the “number 

of leaves” and this is divided into three states 

(none, one, and more than one). In the genera 

Cattleya (Epidendreae) and Spiranthes (Cranichideae) two or more states are known to occur; 

this is also the case with several other genera in 

his study. In Lavarack’s character of the leaf 

bases, one is not sure what he means, but within 

the genus Spiranthes (Spiranthoideae) several different types of leaf base occur. Another of Lavarack’s characters, ovate versus linear leaves, 

has both states often found in the same genus. 

Still another example, the length of the column, 

is so variable that it cannot be used on the subtribal level (e.g., Spiranthinae, Balogh, 1982). 

We could go on, but the point is obvious. By 

looking at only one individual or species one can 

make a judgement; the information that the character varies is either ignored or considered to be 

unimportant. 

Because phenetic studies treat all characters 

equally, they group by plesiomorphies as well as 

apomorphies (see discussion of traditional orchid 

systematics). Character weighting occurs, however, because of the various ways of coding multistate characters. Also, because there is no need 

to thoroughly examine the distribution and morphology of the characters in order to decide on 

homology and polarity, some characters are 

really groups of characters. 

Phenetics has the additional problem of dividing characters into “character states,” which are 

then treated as distinct entities rather than sets 

and subsets. An example from orchids is the 

number of chambers in the ovary. Figure 2 shows 

the difference between treating characters as 

having states and treating them as internested. 

In the Orchidaceae the ovary can be either threechambered or one-chambered. In a phenetic 

study these are treated as two distinct states of 

the same character and both can be used to group 

taxa. So, the use of “character states” is tied 

directly to the procedure of grouping with plesiomorphic characters because when “character 

states” are discrete entities they can be given 

equal value in grouping. Cladistically, the oneand three-chambered ovaries are treated as 

nested. This is easiest to see using a Venn diagram (Figure 2). The three-chambered ovary 

character is plesiomorphic and the one-chambered is apomorhic (evolutionarily novel). The 

one-chambered ovary is treated as a subset of the 

three-chambered one and all taxa that have the 

apomorphy are also considered to have the threechambered ovary because one has developed 

from the other. After the cladogram was completed it was best to hypothesize that the devel- 
NUMBER 61 9 

opment of the one-chambered ovary had occurred twice in the Orchidaceae. 

In addition to the procedural difficulties, we 

find the results of phenetic studies irritating because we are presented with a diagram, but have 

no way of knowing which characters are responsible for the various branches (this is also a problem with some intuitive trees). 

Interestingly the written classification from 

Lavarack’s study does not strictly follow the 

groups indicated in the dendrograms. Rather, it 

shifts genera around when the results are undesirable. In Lavarack’s own words (1976, p. 293): 

This classification does not follow exactly the results presented in Figures 1 and 2 but is a composite of those results, 

the results of other analysis only briefly discussed here, and 

the personal ideas and prejudices of the author. 

It all sounds rather like traditional intuitive systematics. It seems then, that as applied in the 

Orchidaceae, phenetics has not only many of the 

problems of traditional systematics but a few 

additional problems as well. 

Cladistic Analysis of the Orchidaceae 

Character Selection 

Decisions on polarity of characters are based 

on the outgroup comparison method (Hennig, 

1966; Platnick, 1979; Watrous and Wheeler, 

198 l), in which the more general characters are 

plesiomorphic. The character designated by the 

lowest number (1) is the most plesiomorphic one 

for each transformation series (Table 2). 

All characters, both floral and vegetative, were 

considered in this study. Only characters that 

proved to be consistent were used in the cladistic 

analysis, however. Consistent characters are 

those that do not vary within the study taxa. For 

our purposes that meant that the characters had 

to be consistent at the tribal level. Exceptions at 

the species level are not treated in this cladistic 

analysis and are only mentioned under the subtribal descriptions. We consider these exceptions 

to be apomorphies at the species or generic level. 

Some of the characters were consistent in all of 

the tribes except for one or two. The tribes 

where the character was inconsistent were then 

broken down to subtribes for this analysis. The 

only exceptions to this were in the size of the 

pollen unit, i.e., monad or tetrad transformation 

(series 36), and in the cobra-hood-like inner lateral staminode development (character 24-9). 

The presence of tetrads varied so much in the 

Neottioideae that it could not be used even at 

the subtribal level. It was consistent for the other 

tribes, however, so we retained it in the analysis 

and simply listed it as variable for the Neottioideae. Some Epidendroideae have the inner lateral staminodes completely fused to the column 

structure (character 24-4) and some have them 

fused to the column margins and cobra-hoodlike above the anther (character 24-9). While 

these characters were consistent on the genus 

level they were variable on the tribal level and so 

are listed as both on the cladogram (Figure 1, 

Area E). 

The following transformation series are discussed in the order of their occurrence on the 

anther and column (for a listing of the characters 

by number see Table 2). In the remainder of this 

paper, unless otherwise specified, all subfamilies, 

tribes and subtribes are those recognized by 

Burns-Balogh and Funk; a synopsis of these appears at the end of this section (see “Classification 

of the Orchidaceae” p. 60, for a complete listing). 

The 68 apomorphic .characters are grouped into 

42 transformation series ( 14 transformation series contain more than one apomorphic character). The characters fall into three groups, those 

dealing with the pistil, with the stamen(s), and 

with the pollinia. Only a brief description of the 

characters is possible in this paper. A more de- 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

NEU APO CYP SPI NEO 

36( 1-21 

8-3 
NUMBER 61 

ORC 

11 

EPI 

VAN 1 

n 

-‘ 

n 

GAS 

FIGURE 1.-Cladograin of the Orchidaceae. Lines = synapomorphies, closed circles = homoplasies (parallel or convergent evolution), x = loss of a synapomorphy (reversal). 

NEU = Neuwiedioideae; APO = Apostasioideae; CYP = 

Cypripedioideae; SPI = Spiranthoideae, CRA = Cranichideae, Go = Goodyerinae, Tro = Tropidinae, PRA = Prasophylleae, DIU = Diurideae; NEO = Keottioideae, NEO = 

Neottieae, Lis = Listerinae, Lirn = Limodorinae, GEO = 

Geoblasteae, Cal = Caladeniinae, Chl = Chloraeinae, PTE = 

Pterostylideae, THE = Thelyniitreae; ORC = Orchidoideae, 

EPI PLE 

C 

DIS = Diseae, SAT = Satyrieae, ORC = Orchideae; EPI = 

Epidendroideae, GAS = Gastrodieae, TRI = Triphoreae, ARE 

= Arethuseae, VAN = l’anilleae, EPI = Epidendreae, PLE = 

Pleurothaks group, DEN = Dendrobieae, MAL = Malaxideae, 

COE = Coelogyneae, Coe = Coelogyninae, Cor = Corallorhizinae, MAX = Maxillarieae, VAD = Vandeae. Data matrix 

for this cladogram is in Table 3. Constancy index = 60%. 

Numbers listed are found in Table 2. A-F = areas 

of the cladogram referred to in the “Discussion” (see 

page 41). 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE ‘,.--Characters and their respective transformation series used in the “Cladistic 

Analvsis of the Orchidacae.” 

Ti-ansformation series 

Pistil 

1, Fusion of style 

2. Bending of style 

3. Ovary chambers 

4. Fruit type 

3. Endospern1 

6. Stigma modification 

7. \‘iscidium attachment on 

pollinia 

8, Rostel I uiii (modified 

portion of median stigma 

lobe) shape 

9. Stigma 

Character 

0. free 

1. style fused w/filanients 

at bases 

2. style fused w/filanients 

and staminode bases 

3. style fused .-/ 

filaments, staminodes, 

and stigma to column 

4. style extremely 

reduced or absent 

5. style fused w/filament 

and staminode bases, 

style reduced 

1. erect or bending forward 

2. bending backward 

1. 3-chambered 

2. 1-chambered 

1. capsule 

2. berry 

1. present 

2. absent 

1. none or slightly asymmetric 

2. all lobes facing toward 

center of flower 

1. no Liscidium 

2. at or near apex 

3. at or near base 

1. all three stigma lobes 

identical or slightly 

asymmetrical; no ros. 

t el I uin 

2. rostelluiii strap-like, between bases of anther 

theca 

3. rostellum highly reduced 

4. rostellum curled over 

and as long as )vide ot 

wider 

5. rostellum longer thar 

wide 

1. slightly convex 

2. concave 

3. very oblong, extending 

‘A-% of the way dowr 

col u ni n 

10. Rostelluni 

1 1. Clinandrium fusion 

12. Haniulus stipe 

13. Tegular stipe 

;tam en 

14. Number 

Median Outer Stamen 

15. Fusion of entire filament 

only Mith staminodes 

16. Outer median staniinode 

1 f. Anther base position (insertion on filament) 

18. Fusion of outer median 

anther base to column 

apex 

19. Developnient of outermedian stamen in relation to column axis 

20. Locules 

2 1. Anther condition 

2 2. Anther modification 

Character 

1. not sensitive 

2. sensitive 

1. incomplete 

2. complete 

1. absent 

2. present 

1. absent 

2. present 

1. one outer median and 

2. outer median 

3. inner laterals 

two inner laterals 

1. no 

2. yes 

1. none (still an anther) 

2. well developed 

3. filament-like 

1. below stigma 

2. at or near stigma base, 

anther rarely extending beyond rostelluni 

3. at or near stigma apex 

(lateral lobes), anther 

extending above rostelluni 

4. above stigma apex (lateral lobes), anther extending above rostelluni 

1. free 

2. fused 

1. erect to suberect 

2. up-side-down 

3. at right angles to column axis 

4. operculate, incumbent 

to hyperincurnbent 

1. rarely inore than 4 

2. 2 to 12 

1. never deciduous 

2. deciduous (not always) 

1. lily-like 

2. cap-like 

3. bases of thecae usually 

separated, often widely 
NUMBER 61 13 

TABLE P.--Contanued. 

Transformation series ~~ ~ 

Inner Lateral Stamens 

23. Anther tube (adherent 

margins of stamens and 

if present, margins of 

staminode) surrounding 

style 

24. Inner lateral staminode 

development 

25. Auricles 

26. Clinandrium coniposition 

27. Staminode to staminode 

fusion 

Character 

1. absent 

2. present 

1. none (still anthers) 

2. fused to style for most 

of length but apices 

free and projecting, often inconspicuous 

3. long membranous 

4. not apparent, completely fused to column 

structure 

5. fused to margins of column and flared at apex, 

cobra-hood-like below 

anther 

6. fused to margins of column, broad and hooklike at upper %-% 

7. hood-like surrounding 

stigma and anther, 

with pseudostamens or 

other ornamentations 

8. long and thick 

9. fused to column margins and cobra-hoodlike above anther, margins of staminodes 

fused to each other at 

apex and behind or 

above anther 

1. absent 

2. present 

1. absent 

2. back of stigma and 

staminode margins 

3. back of rostellum and 

probably staminode 

margins 

but fused to column at 

some point, sometimes 

only at the very apex 

1. free from each other 

Transformation series 

ollinia 

28. Caudicle composition 

29. Caudicle length 

30. Caudicle to caudicle fu31. Pollinia position 

sion 

32. Pollinia shape 

33. Pollinium set size 

34. Pollinium fusion 

35. Pollen aggregation 

36. Pollen unit 

37. Elastoviscin 

38. Cohesion strands 

39. Aperture 

40. Foot layer 

41. Incipient columellae 

42. Operculate culpus 

Character 

2. inner margins fused 

forming a hood-like 

structure 

1. none 

2. embedded with pollen 

3. sterile; just elastoviscin 

1. short, reduced, con2. tail-like, elongate 

1. free 

2. fused 

1. side by side 

2. superposed 

1. unformed 

2. attenuate apex, 

rounded base 

3. rounded apex, attenuate base 

4. rounded apex and base 

1. each member +/- 

equal 

2. unequal 

1. free 

2. fused to each other in 

set 

1. powdery, non-aggregated, or sticky 

2. granulate, consolidated 

3. sectile 

4. hard, waxy 

1. monad 

2. tetrad 

1. absent 

2. present 

1. absent 

2. present 

1. colpate 

2. porate/ulcerate 

1. present 

2. absent 

1. absent 

2. present 

1. absent 

2. present 

nective 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Characters as States 

Characters as part of a 

Transformation Series 

\ 

FIGURE 2.-Characters as character states and as members 

of a transformation series. This example treats single-chambered vs. 3-chambered ovaries. 

tailed discussion of the characters used in this 

analysis can be found in a series of papers published in the Orchid Review (Burns-Balogh, 1983a, 

b; Burns-Balogh and Funk, 1983) and The Canadian Orchid Journal (Burns-Balogh, 1984a, b; 

Burns-Balogh, Borg-Karlson, and Kullenberg, 

1985). We are not prepared to say which family 

is most closely related to the Orchidaceae but it 

is undoubtably something in the Liliales, so for 

most characters we used the entire subclass as 

the outgroup in this study. The Appendix contains a list of all specimens examined and Table 

3 has the data matrix used in this analysis. 

SYNOPSIS OF BURNS-BALOGH AND FUNK 

CLASSIFICATION 

NEUWIEDIOIDEAE 

APOSTASIOIDEAE 

CYPRIPEDIOIDEAE 

SPIR ANTHOIDEAE 

DIURIDEAE 

PRASOPHYLLEAE 

CRANICHIDEAE: Cranichidinae, Spiranthinae, Manniellinae, Cryptostylidinae, Pachyplectroninae, Tropidiinae, 

Goodyerinae* 

NEOTTIOIDEAE 

GEOBLASTEAE: Caladeninae, Chloraeinae* 

PTEROSTYLIDEAE 

THELYMITREAE 

NEOTTIEAE: Listerinae, Limodorinae* (including Rhizanthellinae) 

ORCHIDOIDEAE 

ORCHIDEAE: Orchidinae, Habenariinae, Huttonaeinae 

DISEAE: Diseae, Coryciinae 

SATYRIEAE 

GASTRODIEAE: Gastrodiinae, Epipogiinae, Stereosandrinae(i), Wullschlaegeliinae(?), Nervilinae 

TRIPHOREAE*: Tropidieae, Acianthinae(?) 

ARETHUSEAE 

VANILLEAE: Vanilliinae, Pogoniinae, Palmorchidinae, LeDENDROBIEAE*: Thuniinae, Glomerinae, Dendrobiinae, 

Pleurothallis Group: Pleurothallidinae, Meiracyllinae, 

EPIDENDREAE: Laeliinae, Bletiinae, Sobralinae 

MALAXIDEAE* 

COELOGYNEAE: Corallorhizinae*, Coelogyninae 

MAXILLARIEAE*: Zygopetalinae, Bifrenariinae, Lycastinae, Maxillariinae, Dichaeinae, Telipoginae, Ornithocephalinae, Polystachyinae, Cryptarrheninae, Calypsoinae 

VANDEAE: Sarcanthinae, Aerangidinae, Angraecinae, Eulophiinae, Cyrtopodiinae, Genyorchidinae, Acriopsidinae, Catasetinae, Stanhopaeinae, Thecostelinae, Oncidiinae, Pachyphyllinae 

EPIDENDROIDEAE 

canorchidinae 

Bulbophyllinae, Sunipiinae 

Thelasinae, Podochilinae, Eriinae, Adrorhizinae 

* Not indicated to be monophyletic groups. 

FIGURES 3-36.-Column morphology of the Orchidaceae. Key to letter symbols: A = anther, S = stigma, 

ST = staminode, V = viscidium, R = rostellum, C = 

column, AU = auricle, CL = clinandrium. Line length 

= 5 mm except in Figure 9, where line length = 1 

mm. 
NUMBER 61 15 

Discussion of Characters and Their 

Aggregation into 

Transformation Series 

CHARACTERS OF THE PISTIL 

In many orchids, most of the bulk of the 

column structure is composed of the style. In 

transformation series 1 (t.s. l), the fusion of the 

style with the various structures results in the 

formation of the column. The basic condition is 

found in Neuwiedioideae (Figure 3), which is 

most similar to other Lilidae with only basal 

fusion of style and filaments, while the diandrous 

orchids (Figures 4-6) have a “column” formed 

by the fusion of the filaments, staminode base, 

and style (Plate 3a). The monandrous orchids, 

on the other hand, have a complete column 

structure (Figures 7-36) resulting from the fuFIGURE 3.-Neuwiedioideae, Neuwiedia singapureana 

(Baker) Rolfe. Booea 25570 (US). 

sion of the style, stigma, staminodes, the filament 

of the fertile anther, and in some cases, i.e., the 

Orchidoideae, the anther base (Plate 3d-f, h-z, 

k-n, Plate 4b-d,f-i, k). In the Diurideae, reduction in the above structures has resulted in the 

“absence” of a column structure (Figure 10) while 

in Prasophyllinae (Figure 11) the column is very 

reduced and composed of only the bases of the 

filament, style, stigma, and staminodes. The style 

is usually erect or bending forward (Table 2, t.s. 

2), but in Diseae (Figure 21) and Satyrieae (Figure 22) the bending at the apex of the column 

results in a reversal in the position of the anther 

(and stigma in Satyrieae). 

Compared to the other families in the monocotyledons the pistil has undergone great modification, including the following: reduction in the 

number of ovary chambers (Table 2, t.s. 3); lack 

of an endosperm (Table 2, t.s. 5); all three or 

two lobes present on one side of the gynoecial 

apex facing the center of the flower (Table 2, t.s. 

6); and the formation of the rostellum from the 

median stigma lobe (Table 2, t.s. 8): The possession of three functional stigmatic lobes is the 

and is therefore considered to be plesiomorphic 

P 

FIGURE 4.-Apostasioideae, Apostasia wallachii R. Brown ex general condition found in the 

Wallich. Dressler s.n. (from photo). 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 3.-Data matrix; number of characters per transformation series in all transformation 

series with more than two characters each apomorphy was treated independently of the others 

and no sequential relationship was inferred until after the cladogram was constructed (B = 

both). 

TAXA/TS I 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 

NEU 1 

NEU 11 

APO 

CYP I 

CYP I1 

Go I 

Go I1 

Tro 

Tro I 

Tro 11 

Tro Ill 

PRA 

DIU 

Lis 

Lim 

Ca I 

Chl 

PTE 

THE 

SAT 

DIS 

ORC 

GAS I 

GAS I1 

TRI 

ARE 

\'AN I 

VAK 11 

EPI 

PLE 

DEK 1 

DEK 11 

hlAL 

Coe 

Cor 

MAX 

\'AD 

11112111111111111111 

11122111111111111111 

21112111111113131111 

21112211111113121111 

21212211111113121111 

31212225112122112111 

31212225112112112111 

31212225112212111111 

31212225112112112111 

31212225112212112111 

31212225112112112111 

51212225111212112111 

41212223111112112111 

31212234121112113111 

31212233111112113111 

31212233111112113111 

31212233111112113111 

31212233111112113111 

31212233111112213111 

32212232111112114221 

32212232111112114231 

31212232211112114211 

31212224211112114111 

31212224211112114111 

31212224211112114111 

31212224212112114111 

31122224212112114111 

31212224212112114111 

31212224212112114112 

31212224212112114112 

31212224212112114111 

31212224212112114111 

31212224212112114111 

31212224212122114111 

31212224212122114111 

31212224212122114111 

31212224212122114111 

in those orchid genera possessing three functional lobes, such as the triandrous and diandrous 

orchids plus isolated monandrous orchids such 

as Cephalanthera and perhaps Rhizanthella. 

The rostellum separates the pollen masses 

from the functionally receptive stigmatic lobes, 

supports the pollinariurn (until removal by the 

pollinator), and provides the sticky material or 

viscidium (by which the pollinia are attached to 

the pollinator). In some genera where a rostellum 

or viscidium is lacking, the stigmatic surface may 

provide the sticky substance. An example of this 

is Cephalanthera. In this paper, the viscidium is 

treated in the broad sense, i.e., any substance 
NUMBER 61 

TABLE 3.-Continued. 

17 

21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 

1111111111111111111112 

1111111111111111111112 

1121111111111111111112 

1111111111111111111221 

1111111111111111111221 

111212131112113 2222111 

111212111112113 2222111 

111212131112113 22 2 2111 

1112121111122122222111 

111212131112112 2222111 

111212111112112 22 22111 

111812131112113 22 2 2111 

1113121111121121222111 

111211111112112B222111 

111211111112112B2 22111 

111511111114112B222111 

111511111113112B2 2 2111 

111611111114112B222111 

111711211113112B222111 

13 11211321131132212211 

1311211321131132212211 

1311211321131132212211 

111411111113113 2212211 

1 1 1 4 1 1 1 3 2 1 1 3 1 1 3 2212211 

111411111113112 2212211 

121913111114113 2212211 

12191 3 1 1 1 1 1 4 1 1 2 2212111 

1219131111141121212111 

22 19131211141142212211 

2219131111121142212211 

121913111114114 2212211 

1219131211141142212211 

1219131111142142212211 

2219131212242142212211 

2219131212242142212211 

2219131311242142212211 

2219131311242242212211 

originating from the rostellum which serves to 

attach the pollen masses to the pollinator. In 

some cases, e.g., Laeliinae, the viscidium is a 

viscid fluid and Dressler (pers. comm., 1982) 

suggests the term “rostellar glue” to distinguish 

it from the solid viscidium that is usually found 

in the orchids. The rostellum assumes a variety 

of shapes and sizes that are represented by the 

five apomorphic characters (Table 2, t.s. 8). 

These are treated independently from one another (Table 4). In those orchids with a welldefined rostellum, the viscidium abscisses from 

the rostellum proper at the time of removal of 

the pollinia and is considered a part of the polli- 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

\ 

FIGURES 5, 6.-Cvpripedioideae, 0)pripedzum acaule Aiton. Balogh 1033 (US). 

nariurn. In some taxa the rostellum is highly 

reduced (Neottioids) and is usually all viscidium. 

Consequently, it is almost totally removed with 

the pollinia. In Listerinae, the rostellum is sensitive to pressure (Table 2, t.s. 10) and exudes a 

viscid droplet when an insect touches it (the 

r oste 11 u m) . 

Other apomorphies of the rostellum include 

the attachment of the viscidium on the pollinia 

(Table 2, t.s. 7). It may be either apical or at or 

near the base of the pollinia. Sometimes a portion 

of the rostellum called a stipe is removed with 

the pollinarium. Rasmussen (1 982) distinguishes 

between the tegular stipe (modified rostellar 

epidermis) of the vandoid orchids and some 

Goodyerinae (Table 2, t.s. 13) and the hamular 

stipe (recurved apex of the rostellum) of the 

Prasophyllinae, some Cranichidinae, and Tropidinae, two anomalous genera, Genoplesium and 

Microtis, now recognized as members of Neottioideae, and at least one species of Bulbophyllum 

of Dendrobieae/Epidendroideae (Table 2, t.s. 

12). Because these two types of stipe are anatomically different we have treated them as different 

(Table 2, t.s. 12, 13; for illustrations of t.s. 7, 12, 

and 13 see section on pollinaria, p. 30). 

The presence of a clinandrium is aponiorphic 

(Table 2, t.s. 11). There are two types: complete, 

lchere the margins of the stigma or rostellum are 

fused to the margins of the staminodes (often 

with a short free apex present) (Figure 8); and 

incomplete, \There there is no fusion betrveen the 

margins of the stigma and staminodes. These two 

apomorphies are treated independently of one 
NUMBER 61 19 

another. 

Other modifications of the pistil include the 

reduction in the number of the ovary chambers 

(Table 2, t.s. 3), a berry fruit (Table 2, t.s. 4), 

and the stigmatic surface that is concave or oblong (Table 2, t.s. 9). The Orchidoideae, Apostasioideae, Neuwiedioideae, two genera in the 

Cypripedioideae, and a few genera in the Vanilleae, have a plesiomorphic three-chambered 

ovary. All other orchids have a single chambered 

ovary. In a few species of Neuwiedia and Vanilleae 

the fruit type is a berry, whereas in all other 

orchids a capsule is found. The stigmatic surface 

is sometimes slightly convex. Because this is the 

condition found in the monocots we have designated it the plesiomorphic character. The apomorphic character of this transition series is the 

concave surface found in some of the Orchidoideae and in Epidendroideae. In Epidendroideae 

the concave stigmatic surface is tucked under the 

roof-like rostellum. This is, in large part, the 

result of the bending of the column. In the 

Pterostylideae the stigmatic surface is oblong and 

extends from approximately 94 to % down the 

column to compose a total of about '/2 of the 

column length (Figure 16). In Habenuria, the 

stigmaphores support the individual stigmatic 

lobes, but we have not included this in our diagrams because we are not including characters 

that are used below the rank of subtribe. 

CHARACTERS OF THE STAMEN 

The stamen is the least modified structure on 

the column (Table 2, ts. 14-27). Each flower in 

the monocotyledons has six stamen positions, 

only three of which ever bear any structures in 

the Orchidaceae. Figure 37 illustrates that only 

the outer median and inner lateral positions are 

occupied in the orchids. The Neuwiedioideae 

have all three stamen positions occupied. In the 

diandrous orchids the inner lateral stamens are 

retained and the outer median one has developed 

into a staminode. The monandrous orchids, on 

the Other hand, have retained Only the outer 

median stamen and both of the inner lateral 

FIGURES 7, 8.-Spiranthoideae, Cranichideae, Spiranthes 

cerlzua (L.). L.C. Rich. Balogh 914 (US). 
C 

I S I 

ST \ 

FIGURE 9.-Spiranthoideae, Cranichideae, Cranichis sp. 

Greenwood 825 (AMO). 

A 

S 

FIGURE 1 0.-Spiranthoideae, Diurideae, Diuris maculata 

Smith. Clements s.n. (Balogh). 

R 

S 

A 

S 

FIGURE 12.--Neottioideae, Neottieae, Listera ouata (L.) R. 

Brown. Kurzweil s.n. (Balogh). 

A 

S 

C 

FIGURE 1 3 (above).-Neottioideae, bieottieae, Cephalanthera austinae (A. Gra?) Heller. Sandberg and Leiberg 571 

(US). 

FIGURE 1 1 (left).-Spiranthoideae, Prasophylleae, Prasophjllum elatum R. Brown. Clernents s.n. (KCBG). 
NUMBER 61 21 

A 

S 

C 

FIGURE 14.-Neottioideae, Neottieae, Epipactis palustrzs (L.) 

Crantz. Kurzweil s.n. (Balogh). 

FIGURE 

LA 

R 

ST 

FIGURE 16.-Neottioideae, Pterostylideae, Pterostylis ~ufu R. 

Brown. Clements s.n. (NCBG). 

5.-Neottioideae, Thelymitreae, Thelymitra carnea 

R. Broivn. Clements 2383 (NCBG). 

FIGURE 17.-Neottioideae, Geoblasteae, Caladenia catenata 

(Smith) Druce. Clements s.n. (NCBG). 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 19.-Orchidoideae, Orchideae, Serapias parvzjlora FIGURE 21 .-Orchidoideae, Diseae, Dim crassicornis 

Parl. Kurzweil s.n. (Balogh). Lindley. Wood 8430 (US). 
NUMBER 61 23 

FIGURE 22.-Orchidoideae, Satyrieae, Satyrzum nepalense 

Don. Kurzweil s.n. (Balogh). 

I 

FIGURE 23.-Epidendroideae, Triphoreae, Triphora trianthrophora (SW) Rydberg. Greenwood 11 50 (AMO). 

FIGURE 24.-Epidendroideae, Vanilleae, Vanilla planifolia 

Andrews. No collector, s.n. (SEL). 

FIGURE 25.-Epidendroideae, Epidendreae, Arpophyllum sp. 

Hagsater 4502 (AMO). 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 26.-Epidendroideae, Epidendreae, 

Greenwood 10 10 (AMO). 

Bletia sp. 

FIGURE 28.-Epidendroideae, Epidendreae, Coelia bella Reichenbach filius. Greenwood 540 (AMO). 

A 

S 

FIGURE 27 .-Epidendroideae, Epidendreae, Hexalectris 

brevicaulis L.O. M’illiams. Greenwood 1059 (AMO). 

FIGURE 29.-Epidendroideae, Epidendreae, Encyclza cochleata (L.) Dressler. Balogh s.n. (Balogh). 
NUMBER 61 25 

‘‘i I 

FIGURE 30.-Epidendroideae, Epidendreae, Sobralia macrantha Lindley. No collector, s.n. (SI). 

stamens have become staminodes. The only exception to this is in Diplandorchis, which has two 

median stamens, and another Chinese genus, 

Tangtsinia, which has five staminodes and a single 

outer median stamen. The possession of all three 

functional stamens is the plesiomorphic condition for transformation series 14 (Table 2). Because the diandrous and monandrous conditions 

involve different stamens, they are treated as 

separate apomorphies and are in no way interdependent. Because no relationship is indicated 

among the staminodes that have developed from 

different stamens, the characters of the stamens 

have been divided into those that deal with the 

outer median stamen (Table 2, t.s. 14-22, Figure 

37) and those that deal with the inner lateral 

stamens (Table 2, t.s. 23-27). 

There are other monocot families that exhibit 

reductions in the number of stamens (Figure 37). 

FIGURE 3 1 .-Epidendroideae, Epidendreae, Calypso bulbosa 

(L.) Oakes. Peterson s.n. (Balogh). 

In the Zingiberaceae, there is a single functional 

anther in the inner median position and four 

staminodes, two in the inner lateral positions and 

two in the outer lateral positions. The fusion of 

the two outer staminodes has produced a “labellum” that has replaced the median petal. The 

Marantaceae also have a single functional anther 

with the outer stamens represented as staminodes. In the Commelinaceae there are three 

fertile adaxial stamens in many taxa. Only two 

families however, have the same type of stamen 

situation as that found in the orchids. In the 

Pontederiaceae, Heteranthera, has a stamina1 situation identical to Neuwiedia, and Hydrothrix has 

a single median stamen as in the monandrous 

orchids. In the Philydraceae a single fertile stamen is in the same position as that found in the 

monandrous orchids (Vermeulen, 1966). Because of the lack of any other similar apomor- 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

A 

R 

S 

FIGURE 32 (left, upper).-Epidendroideae, Coelogyneae, 

Coelogyne mayerianu Reichenbach filius. No collector, s.n. 

(SEL). 

FIGURE 33 (left, lower).-Epidendroideae, Coelogyneae, 

Corallorhiza striata Lindley . Balogh 977 (US). 

FIGURE 34.-Epidendroideae, Vandeae, Gongora truncata 

Lindley. No collector, s.n. (SEL 105-76-25). 

phies, the most parsimonious explanation for 

these stamina1 losses is one of parallel evolution, 

with these characters being autapomorphies for 

the respective groups. (Autapomorphies are of 

no use in determining relationships because they 

found in only one taxon.) 
NUMBER 61 27 

FIGURE 35.-Epidendroideae, Vandeae, Chondrorhyncha 

lendyana Reichenbach filius. No collector, s.n. (SI 79820). 

Outer Median Anther Position 

A unique situation is found in the outer median anther filament of the Thelymitreae. The 

anther filament is almost completely free of the 

style but is fused to and incorporated into the 

staminodal structure (Table 2, t.s. 15; Figure 15). 

The general condition is the fusion of the filaFIGURE 36.-Epidendroideae, Vandeae, Trichoglottis fasciata 

Reichenbach filius. No collector, s.n. (SEL). 

ment to the style and the margins of the staminodes. 

The outer median anther is inserted on the 

filament “at or near, rarely below, the base of 

the stigma” (and parallel to the stigma) (Figures 

7-1 l), “at or near the apex of the stigma” (Figures 12-1 7), or “above the apex of the stigma” 

(Figures 18-36), of which the latter two have an 

anther that extends beyond the stigma apex. 

Because these two insertions are found only in 

some groups of orchids and probably nowhere 

else in the monocots, they are considered to be 

apomorphic (Tables 2 and 4, t.s. 17). Similarly, 

the fusion of the anther base to the column apex 

is found only in the Orchidoideae (Table 2, t.s. 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

LI LI ACE AE NEUWlEDlA 

APOSTASIA 

CY PR IPEDIUM 

MONANDROUS ORCHIDS ZINGIBERACEAE CANNACEAE 

FIGURE 37.--Staminal circles in Liliaceae, Orchidaceae, Neuwiedia, Apostasia, Cyprepedium, 

nionandrous orchids), Zingiberaceae, and Cannaceae (triangles = staminodes, double solid 

circles = anthers). For simplicity all groups are shown in the resupinate condition. 

18; Figures 18-22). In the transformation series 

dealing with the development of the outer median stamen (Table 2, t.s. 19), once again, the 

plesiomorphic character (erect to suberect) is 

found in the monocots, e.g., Liliales. The three 

apomorphic characters-the “upside down” anther (Satyrieae; Figure 22), the anther at right 

angles to the style (Diseae; Figure 21), and the 

operculate incumbent to hyperincumbent anther 

(higher Epidendroideae; Figures 25-36)-are 

treated as independent of one another. 

The majority of orchids have four separate 

anther locules, like other monocots (although it 

is often difficult to see the median partitions 

between the pairs of locules). In some orchids 

such as the genus Caladenia (Geoblasteae), and 

in some Epidendreae and Pleurothallid groups, 

the locules are divided so that there may be 

anywhere from 6 to 12 separate pollen masses 

(for illustrations see discussion on pollinaria, p. 

30). In Pleurothallidinae and isolated Epidendreae, there may be a loss, or perhaps fusion, of 

two locules so that there appear to be only two 

pollinia per pollinarium. There are some genera 

in the Vandeae in which there also appear to be 

only two pollinia, but this is the result of fusion 

and there are still four locules. The variation in 

the number of locule partitions is considered to 

be apomorphic (Table 2, t.s. 20), however, because the numbers of pollinia are not consistent 

at the subtribal level in Laeliinae or Pleurothallidinae, they are treated as single apomorphies. 
NUMBER 61 29 

In many of the orchids, especially in tribes 

Coelogyneae, Maxillarieae, Vandeae, and Epidendreae (sensu Burns-Balogh and Funk) the 

filament breaks at the juncture of the anther 

when the pollinarium is removed, causing the 

anther to fall off the column (Table 2, t.s. 21). 

The anther can be either lily-like (plesiomorphic, Figure 3), cap-like (Figures 25-36), or 

with the connective separating the anther halves 

at their bases (rarely completely (Figures 18-22; 

Table 2, t.s. 22). A peculiar situation has been 

found in two species of Goodyera, i.e., G. repens, 

and G. pubescens. The anther appears to be similar to that of some Epidendroideae in being 

nearly incumbent and cap-like on the column 

apex. There is no doubt that these two closely 

related species have a specialized anther. 

In the diandrous orchids the outer median 

stamen has developed into a staminode that is 

either filament-like (Figure 4) or well developed 

(Figures 5-6). The two situations are treated as 

independent apomorphies (Table 2, t.s. 16). 

Inner Lateral Stamen Positions 

There are fewer apomorphic characters concerning the inner lateral stamens/staminodes. 

Twelve apomorphies are organized into four 

transformation series (Table 2, t.s. 24-27) and 

one transformation series is concerned with the 

adherence of the inner lateral stamen margin 

(Table 2, t.s. 23). In the Apostasioideae the margins of the stamens and, if present, the margins 

of the staminode, are adherent marginally into a 

tube-like structure that surrounds the style for a 

portion of its length (Figure 4). There is variation 

in the length of the style in the species of Apostasia (de Vogel, 1969). It may be that the style 

length varies according to age of the plant and 

that the anthers release their pollen onto the 

style surface as it grows up through the anther 

tube (H. Robinson, pers. comm., 1983). This 

character (Table 2, t.s. 23) is not present elsewhere in the monocots or the orchids. Consequently, it is considered to be apomorphic. 

The development of the staminodes and how 

they are fused with the style is variable (Table 2, 

t.s. 24). Unfortunately, the distribution of staminodal characters in the Epidendroideae is not 

well known so that we could not plot the exact 

distribution for the various types in this subfamily. Each of the known apomorphies are treated 

as independent of one another (Table 4). There 

is some question as to whether the auricles found 

in the Orchidoideae (Table 2, t.s. 25; Figures 

18-22) are staminodes or a different structure. 

Vermeulen (1 966) thinks that because there are 

no vascular traces in the auricles and because 

when supernumerary stamens develop these 

structures are also present, that the auricles are 

not staminodes. Others, such as Schlechter 

(1 970-1 984) think auricles are staminodes because they occur in the same position as other 

staminodes. Whether they are staminodes or not, 

these structures are easily designated as apomorphies because they have only scattered occurrence elsewhere in the monocots. Dressler (pers. 

comm., 1983) reports that Vermeulen thinks 

they also exist in Allium while he (Dressler) believes Lyperanthus (Orchidaceae) has auricle-like 

staminodes. If they are treated as staminodes 

there would be another independent apomorphy 

in t.s. 24 (Table 2) and t.s. 25 (Table 2) would 

be deleted. 

The clinandrium (Table 2, t.s. 26) is present 

in two subfamilies, Spiranthoideae and Epidendroideae. In the Spiranthoideae the clinandrium 

is composed of the back of the stigma/rostellum 

and the margins of the staminode apices. In the 

Epidendroideae the clinandrium is composed of 

the back of the rostellum and probably the margins of the staminode apices. 

The final stamen transformation series is concerned with staminode to staminode fusion (Table 2, t.s. 27). In almost all orchids the staminode 

margins are fused to the column, but in the 

Thelymitreae the staminode apices are free of 

any pistil structure and instead are fused to one 

another (and the filament) to form a hood-like 

structure over the stigma. In addition the staminodal hood has various ornamentations on it, 

resembling, for instance, what appear to be stamens or other pollinator attracting devices. 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

CHARACTERS OF THE POLLINARIUM 

Pollinarium structure has been used to circumscribe large groups of orchids (Lindley, 1830- 

1840; Richard, 1817) and continues to be the 

source of important characters in the family. The 

tradition has been maintained to some extent in 

this classification as most major groups are defined by at least one apomorphy of the pollinarium. 

In the monandrous orchids the pollen masses 

and any accessory structures take on a characteristic shape called the pollinarium. The shape of 

this structure is usually diagnostic to genus 

(rarely to species). Naturally, a powdery, unstructured pollen mass or pollinium is the plesiomorphic condition. There are three apomorphic 

types of pollen aggregation (Table 2, t.s. 35): 

granulate, composed of monads or tetrads (Figures 38-42, 48-50, 52, 57-62); sectile or massulate, composed of packets of tetrads, rarely 

monads (Figures 43-47, 51, 53-56, 63-65); and 

pollinia, hard and waxy, composed of compacted 

tetrads that share a common wall (Figures 66- 

84). Each of these was treated as an apomorphy 

independent of the other. A very rare condition 

occurs in Chiloglottis (Geoblasteae), which has 

sectile pollinia composed of monads (Figures 5 1, 

53) or tetrads (Ackerman and Williams, 1981). 

The majority of orchids have some type of 

accessory structure accompanying the pollinia 

and viscidium (Table 2, t.s. 28-30). These structures can be pollen-embedded caudicles or sterile 

caudicles with a stipe (Table 2, t.s. 28). Some 

members of the Spiranthoideae and the Neottioideae have the simplest type of pollinarium, 

i.e., four pollinia and a viscidium (Figures 38- 

42). In the Orchidoideae a tail-like sterile caudicle (Table 2, t.s. 29) and viscidium accompany 

the sectile pollinia (Figures 54-56). All of the 

above types are found in Epidendroideae in addition to fused pollen-embedded caudicles (Table 

2, t.s. 30; Figures 78-79) and extremely reduced 

pollen-embedded caudicles (Figure 7 1). The 

Dendrobieae and Malaxideae (Figures 72-73) 

lack any specialized structures (except for one 

species of Bulbophyllum that has a hamular stipe). 

All apomorphies were treated as independent of 

one another (Table 2). 

In most orchids the pollinia have a definite 

polarity with the apex attenuate and the base 

rounded, as in the Spiranthoideae (Figures 38- 

47) and some Epidendroideae (Figures 68-70). 

In the Orchidoideae and Neottioideae the pollinia have a rounded apex and attenuate, sometimes rounded base, the result of the basitonic 

attachment of the viscidium (Figures 48, 49, 51, 

52, 54, 55). In most Epidendroideae the pollinia 

are rounded at both the apex and base (Figures 

56, 7 1-84). All three conditions are considered 

apomorphies and are treated independently (Table 2, t.s. 32). 

Elastoviscin (Table 2, t.s. 37) is a male sporogenous breakdown product that is probably present in all monandrous orchids in some form 

(Rasmussen, 1983; pers. comm., 1983; pers. observ. by Burns-Balogh). In the Spiranthoideae 

(Figure 46) and the Neottioideae (Figure 53) the 

elastoviscin is elusive in nature. For example, in 

Ludisia discolor elastoviscin may or may not be 

present (Figure 46) and in Sarcoglottis some species may or may not have degenerated pollen 

tetrads (elastoviscin) at the very apex of the pollinia. It may also take the form of a thin sheathlike layer around the pollinia. In these two 

subfamilies the presence of elastoviscin is considered to be too variable to use as a character. In 

some Spiranthoideae, Orchidoideae and Epidendroideae the elastoviscin takes the form of a taillike or connective caudicle (mentioned above). 

The pollen unit in subfamilies Neuwiedioideae, Apostasioideae, and Cypripedoideae are 

monocolpate monads (Table 2, t.s. 36 and 39), 

but in the monandrous orchids the pollen unit is 

usually a monoporate tetrad. In subfamily Neottioideae and some lower Epidendroideae the pollen unit can be either a monoporate monad or 

chidaceae. Key to letter symbols: V = viscidium, H = 

hamular stipe, P = pollinia, E = elastoviscin, M = 

massular, S = tetrad sheath, PO = pore, C = caudicle, 

T = tegular stipe. 
NUMBER 61 31 

FIGURES 38-42.-Spiranthoideae: 38, Cryptostylidinae, (;ryptosty/is subulata (Labill) H.<;. Reichenbach filius. Bernhardt s.n. (Balogh), X 20; 39, Spirdnthinae, Spiranthes lacera (Raffinesque) 

Kaffinesque. Balogh 904 (US), X 30; 40, 4 1, Cranichidinae, Solenocentrum costaricense Schlechter. Dressler 5841 (Balogh), X 50, X 150; 42, Diurideae, Diurzs maculata Smith. Bernhardt 

s.ll. (Balogh), x 25. 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 43-47.-Spiranthoideae: 43, Prasophplleae, Prasophyllum suttonii R. Rogers and B. 

Rees. J.T. 1340 (SEL), X 40; 44, Tropidinae, Corymborkisforcipigera (Reichenbach filius and 

Warcs.) Williams. Turckheim 860 (US), X 40; 45-47, Goodperinae: 45, Goodyera oblongqolia 

Rafinesque. Balogh I055 (US), X 30; 46, Ludisia discolor (Ker-Gawl.) Richards. Balogh s.n. 

(Balogh), X 50; 47, Physurus sagreanus Richards. Jimenez 2757 (US), X 150. 
NUMBER 61 33 

FIGURES 48%53.-Neottioideae: 48, Pterostylideae, Pterostylis baptisii Fitzgerald. Bernhardt s.n. 

(Balogh), X 90; 49, Thelymitreae, Thelymitra carnea R. Brown. Clements 2383 (NCBG), X 30; 

50, Neottieae, Epipactis gigantea Douglas ex Hooker. Hall 2370 (US), X 2000; 51-53, Geoblasteae: 5 1, 53, Chiloglottis gunnii Lindley, Bernhardt s.n. (Balogh), X 50, X 750; 52, Caladenia 

(carnea) catenata (Smith) Druce. Bernhardt s.n. (Balogh), X 60. 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 54-56.-Orchidoideae: 54, Satyrieae, Satyrium corriJolzum SW. Clements s.n. (KCBG), 

X 40; 55, Diseae, Disa crassicornis Lindley, Wood 8430 (US), X 50; 56, Orchideae, Habenuria 

sp., Balogh 727 (US), X 35. 
NUMBER 61 

FIGURES 57-62.-Epidendroideae, Vanilleae: 57, Vanilla pauczyora Dressler, Dressler s.n. 

(Balogh), X 3500; 58, Palmorchis sp., Dressler s.n. (Balogh), X 100; 59, Lecanorchis brachycarpa 

Ohwi, Ohiui and Walker 7023 (US), X 3000; 60, 62, Psilochilus sp., Dressler s.n. (Balogh), X 

2500, X 35; 61, Cleistes divaricata (L.) Ames, Chapman s.n. (US), X 500. 

35 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 63-67.-Epidendroideae. 63-65, Gastrodieae: 63, Gastrodia sesamoides R. Brown. 

Bernhardt s.n. (Balogh), X 40; 64, Epipogium aphyllum (Schmidt) SW, Baenitz s.n. (US), X 20; 

65, Arethusa bulbosa L. Cheney, s.n. (US), X 100; 66, 67, Epidendreae: 66, Arpophyllum 

giganteum Hartweg ex Lindley, Suarez 171 (AMO), X 100; 67, Schomburgkia tibicinis Bateman, 

Greenwood s.n. (AMO), X 40. 
NUMBER 61 37 

FIGURES 68-73.-Epidendroideae. 68-70, Pleurothallis group: 68, Octarrhena condensata 

(Ridl.) Holtt., Dressler s.n. (Balogh), X 200; 69, Lepanthes sp., Suarez 294 (AMO), X 100; 70, 

Meiracyllium trinasutum Reichenbach filius. Dressler, s.n. (Balogh), X 55; 7 1-72, Dendrobieae: 

71, Sunipia racemosa (SM) T. Tang and F.T. Wang, Dressler s.n. (Balogh), X 130; 72, 

Dendrobium discolor Lindley, Bernhardt s.n. (Balogh), X 50; 73, Malaxideae, Malaxis marsupichila Upton, Clements 1148 (NCBG), X 200. 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 74-79.-Epidendroideae. 74-77, Maxillarieae: 74, Telipogon klotscheanus Reichenbach filius, Dressler s.n. (Balogh), X 30; 75, Calypso bulbosa (L.) Oakes, Peterson s.n. (Balogh), 

X 40; 76, Cryptarrhena guatemalensis Schlechter, Dressler s.n. (Balogh), X 80; 77, Maxillaria 

sp., Balogh 815 (US), X 30. 78-79, Coelogyneae: 78, Govenia liliacea (Llave and Lex) Lindley, 

Greenwood 556 (AMO), X 55; 79, Coelogyne pandurata Lindley, no collector, s.n. (SI), X 25. 
NUMBER 61 39 

FIGURES 80-84.-Epidendroideae, Vandeae: 80, Gastrochilus bellinus Kuntz, no collector, s.n. 

(Frankfurt Botanical Garden), X 20; 81, Dipodium punctatum (Smith) R. Brown, Bernhardt s.n. 

(Balogh), X 40; 82, Fernandezia hartwegii (Reichenbach filius) Garay and Dunsterville, Dressler 

s.n. (Balogh), X 50; 83, Gongora gratulabunda Reichenbach filius, Mejia s.n. (Balogh), X 20; 

84, Plectrophora alata (Rolfe) Garay, JFM 882 (SEL). 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

tetrad. A specialized condition is found in Lecanorchis and Epistephium (Figure 59) (Ackerman 

and Williams, 1980; Burns-Balogh and Robinson, 1985) and possibly Vanilla (Figure 57) in 

which the monad is polyporate. The tetrad condition is considered to be the apomorphic character because almost all other monocots have 

pollen dispersed as monads. 

There are a number of transformation series 

dealing with the pollinia that have only one apomorphic character and are easily polarized because they are found in restricted groups and no 

where else in the monocots: 

1. In most orchids the pollinia are side by side 

(Table 2, t.s. 31; e.g., Figures 39-41); in the 

vandoid orchids, however, there is a situation 

called superposed pollinia in which the locules 

have moved to the margins of the anther and the 

pollinia in each set have a rounded appearance 

(Figures 80-84). In most Vandeae the pollinia in 

each set are fused (Table 2, t.s. 34) so that the 

sets look like spheres and there appear to be only 

two pollinia (Figures 80-83). In other Vandeae 

the fused sets are flat with a cleft delimiting the 

pollinia in each set (Figure 84). 

2. Of all the orchids, only Spiranthinae and 

Cryptostylidinae (both are Cranichideae) and the 

Vandeae, Maxillarieae, Coelogyneae, and Malaxideae (all are Epidendroideae) have unequal sets 

of pollinia (Table 2, t.s. 33; Figures 38, 39). 

3. The presence of cohesion strands is an apomorphic character (Table 2, t.s. 38) found in 

Spiranthoideae and Neottioideae. Cohesion 

strands are acetolysis-resistant structures and 

should not be confused with viscin strands that 

are performed sporopollenin threads. Unlike viscin strands, cohesion strands only assume their 

irregular shape after the pollen tetrads are pulled 

apart. 

4. The presence of an operculate colpus is an 

apomorphic character (Table 2, t.s. 42) found in 

Neuwiedia (Newton and Williams, 1978; Schill, 

1978) and Apostasia (Schill, 1978). We know of 

no other monocot group that possesses an operculum. 

5. In Cypripedioideae, Orchidoideae, and 

most Epidendroideae the exine foot layer is absent, presumably through loss (Table 2, t.s. 40). 

Although the accompanying characters in each 

group indicate that the loss of the foot layer in 

these subfamilies are three different apomorphies, they were coded as the same because additional information on the pollen wall is still 

needed before we can be sure of their classification. 

6. In the Cypripedioideae the loss of the foot 

layer is accompanied by the presence of incipient 

columellae (Table 2, t.s. 41), which is unique in 

the orchids and perhaps in the monocots (BurnsBalogh, 1983a). 

CHARACTERS NOT USED IN THIS 

CLASSIFICATION 

Exine sculpturing, vegetative characters such 

as leaf-type, root-stem tuberoids, corms, pseudobulbs, and perianth characters were not used in 

this cladistic study because they are inconsistent 

at the tribal level. Most of these characters have 

been used to define tribes or subfamilies in past 

classifications. The consistency of the pollen mass 

has also been used, especially by Lindley (1830- 

1840) and Schlechter (191 1, 1926), but is an 

inconsistent character when used alone without 

the accessory structures. This reasoning also applies to the presence of nectaries and shape and 

size of the column foot. Most of these characters 

(or groups of characters) can be used at the 

generic level or below, but usually not at the 

subtribal, tribal, or subfamily levels. In addition, 

the presence of a labellum and the symmetry of 

the flower were not used as characters in this 

analysis because it was impossible to determine 

whether their presence was apomorphic or plesiomorphic in the Orchidaceae. 

Cladogram Construction 

The cladogram was constructed by parsimoniously arranging taxa so as to create the fewest 

number of character conflicts. The final cladogram was the one that required us to make the 
NUMBER 61 41 

fewest number of ad hoc assumptions of parallel 

or convergent evolution or reversals. This procedure is called parsimony. The cladogram was 

constructed by examining a series of three taxon 

statements. The cladogram should be viewed as 

a hypothesis of relationships and not as “truth” 

or “fact.” Cladograms produced by Hennig’s 

method are direct reflections of the data; should 

the information change sufficiently, the cladogram will also change. 

Discussion of Characters as Displayed 

on the Cladogram 

Because the transformation series that contained more than one apomorphic character 

were coded in a conservative manner (apomorphic characters were treated as independent 

occurrences) we can now examine these apomorphies on the cladogram (Figure 1) and see if an 

alternative order is indicated (Table 4). One must 

be careful to treat these statements on the relationship of the apomorphies within the transformation series as hypotheses subject to change if 

and when additional information becomes available. In the following discussion of transformation series refer to Table 4. 

T.S. 1.-All orchids have some fusion of the 

style and filaments. The partial fusion of the 

Neuwiedioid filaments and style can be considered independent of the anther fusion type or as 

a separate event. We have chosen to display it as 

one fusion series rather than have the initial 

fusion occur twice. From this partial fusion (1-1), 

first the style, filament, and staminode bases 

fused (1 -2), then the style, filament, staminodes, 

and stigma fused to form the column (1-3, Area 

A). The two remaining apomorphies for this 

transformation series developed from the fused 

style/stigma/filament/staminodes. The transformation series has therefore been drawn to reflect 

this change. 

T.S. 7.-The basal viscidium attachment can 

be interpreted as either developing once or twice 

from the apical attachment (Table 4). Both interpretations are equally parsimonious and give the 

same branching pattern. Although the former 

interpretation is the one indicated on the cladogram, we have no preference (Table 4). 

T.S. 8.-The cladogram shows that the evolution of the rostellum is more complex than was 

originally coded (Table 4). Rather than each 

apomorphy being derived independently from 

three equally sized stigma lobes, the most parsimonious explanation is as follows: the apical 

stigma lobe developed in five independent directions, the strap-like rostellum of the Orchidoideae (8-2), the curled over one of the Epidendroideae (8-4), and the long rostellum of the 

Spiranthoideae (minus the Diurideae, 8-5). 

There was a parallel reduction of the rostellum 

(8-3) in the Diurideae (Spiranthoideae) and in 

the Neottioideae followed by a reversal to a 

rostellum longer than wide (8-5) in the Listerinae. 

T.S. 14.-As was expected, character 14-2 is 

a synapomorphy for the monandrous orchids 

(Area A) and the Apostasioideae and Cypripedioideae are united by the synapomorphy of 

having only inner lateral stamens (14-3). The 

Neuwiedioideae, which have the plesiomorphic 

character of three stamens (14-1), are left out of 

either group. 

T.S. 17.-The change in anther insertion on 

the filament could be sequential (Table 4) in that 

it progresses from inserted below the stigma (1 7- 

1) to at or near the stigma base (1 7-2; Area A) in 

the monandrous orchids, to at or near the stigma 

apex (17-3; Area B) and finally to above the 

stigma apex (17-4; Area C). In apomorphy 17-2 

the anther parallels the stigma and in characters 

17-3 and 17-4 the anther extends beyond the 

stigma apex. It would not change the branching 

sequence to interpret the apomorphies as independently derived, however, it would require 

repeated fusion of the same structures, so we 

prefer the sequential order. 

T.S. 24.--When present, the inner lateral 

staminodes have one of eight apomorphies. The 

apomorphies were coded as independent and the 

cladogram supports this with the exception of 

inconspicuous staminodes (24-2) and the re- 
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 4.-Character relationships within selected transformation series as they were coded 

and after the cladogram was constructed. 

Transformation After Cladogram 

Series Number As Coded Construction 

1 

7 

a 

13 

17 

5 

t 5 

3- 2 t 

34 t 3-2 

2 \? ?/ 5 4 or t 

16 0 t 4 

2,, /3 

1 

4 3 

\7 

2-1 -5 

2 

t 

1 

1 

t 

0 

f2 

7 r” 3 

2 or 

t 

1 

1 

I 

2 

t 

2+1 

5 

t 

3 

t 

1-0 

2\ f2 

t 

3 

t 

2 

1 

2 

or f 

2- 142 

1 

3 t 3 

t or t 2 

1-2 261 44 t 

4 

t 

3+ 1-4 t 

1 
NUMBER 61 43 

TABLE 4.-Continued. 

Transformation After Cladogram 

Series Number As Coded Construction 

24 

28 

32 

56 

1\1 f!? 

4 

3-1-8 

9 

J1 

2 

2+ 1-3 

3 

t 

261-4 

3 

t 

35 2- 1-4 

9 9 

2 c- 1'-7 

2 I8 

1 

t 

23 

2 

t 

t 

2\r f3 

4 

3 

26 1 

33 

77 r/3 

36 2 -4 

t 

1 

peated development of stamens fused to the col- Sterile caudicles also characterize the Orchidoiumn margins and cobra-hood-like above the an- deae and many of the Spiranthoideae (minus the 

ther (24-9) from a completely fused column Diurideae). This is one of the most variable transstructure (24-4). formation series on the cladogram. 

T.S. 28.-In the Epidendroideae caudicles T.S. 32.-The pollinia shape can be interembedded with pollen (28-2) have occurred preted as twice having developed a rounded base 

three times and sterile caudicles (28-3) twice. (32-2) and twice having developed a rounded 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Qoo Goo Tro I II IH PRA DNJ Goo 000 Tro I n M PRA Dlll 

FIGL~RES 85-8i.-C:ladogr;iiiis sho\\.ing three possible grotipings of tam \\.ittiin the siibfaiiiil~ 

Spiranthoideae (onl) the characters important to this grouping question are illustrated-see 

Figure 1 and Table 3): 85, contains the hypothesis of homoplasy for character 35-3 (12 steps); 

86, contains the hypothesis of reversal for character 3.53 (1 2 steps); 87, contains the hypotheses 

of hoiiioplasy for characters 1 1 and 24-2 (1 3 steps) (Goo = Goodyeriana). 

apex (32-3) from the primitive condition of both 

base and apex being attenuate. The rounded 

apex and base (32-4) developed from the roundattenuate condition once and from this the apex 

became rounded once again (Table 4). 

T.S. 35.-Sectile pollen aggregation has developed five times from the granulate condition: 

(1) Orchidoideae, (2) Arethuseae (Epidendroideae), (3) Gastrodieae (Epidendroideae), (4) 

Goodyerinae-Tropidinae (Spiranthoideae), and 

(5) Prasophyllinae (Spiranthoideae), which makes 

this one of the most variable characters in this 

study. 

Five multi-apomorphy transformation series 

were found to remain as they were originally 

coded. These are t.s. 14, 16, 19, 22, and 26. 

Several transformation series with only one 

apomorphy showed some homoplasy (Figure 1). 

The one-chambered fruit (Table 2, t.s. 3; Figure 

1) showed up in part of the Cypripedioideae and 

at Area A. The berry fruit (Table 2, t.s. 4; Figure 

1) is in some Neuwiedioideae and in some Vanilleae. Stigma modification (Table 2, t.s. 6; Figure 1) is present in all of the Cypripedioideae 

and at Area A (monandrous orchids). The hamulus (Table 2, t.s. 12) appears three times in the 

Spiranthoideae and the tegulus (Table 2, t.s. 13) 

once in the Spiranthoideae and once in the Epidendroideae with a reversal in the Coelogyneae 

(Table 4). It is equally parsimonious to hypothesize that the tegular stipe developed three times 

but this assumption does not alter the branching 

sequence of the cladogram. Deciduous anthers 

(Table 2, t.s. 21; Figure 1) appear twice in the 

Epidendroideae. Tail-like caudicles (Table 2, t.s. 

29; Figure 1) are characteristic of the Orchidoideae and some species of the Gastrodieae (Epidendroideae). Unequal pollinium sets (Table 2, 

t.s. 33) are found in two widely separated groups, 

the higher Epidendroideae and some species of 

the Cranichideae (Spiranthoideae). All monandrous orchids (Areas A-F of cladogram, Figure 

1) have tetrad pollen units (Table 2, t.s. 36) 

except for some species of the Vanilleae (Epidendroideae), and in the taxa of the Neottioideae 

where it varies from monad to tetrad in every 

group. So variable is it in the Neottioideae that 

it is simply listed on the cladogram as 36-1/2. 

Cohesion strands (t.s. 38) are in both the Spiranthoideae and Neottioideae (see “Discussion of the 
NUMBER 61 45 

Cladogram”). The lack of a foot layer (t.s. 40) 

occurs twice, the Cypripedioideae and Area C, 

but it reverses in the Vanilleae. The operculate 

colpus (t.s. 42) is found in the Neuwiedioideae 

and maybe Apostasioideae. 

Transformation series with only one apomorphy that showed no homoplasy or reversals are 

as follows: 2, 5, 10, 15, 18, 20, 25, 30, 31, 34, 

37, 39, and 41. 

Based on the information gained by studying 

the cladogram the multi-apomorphic transformation series have been drawn to eliminate unnecessary reversals. 

The characters that are shown to have evolved 

in parallel could be recoded as different characters. We have found them useful, however, when 

studying the pollination and ecology of various 

groups of orchids and prefer to leave them on 

the cladogram and merely indicate that they 

occur more than once by using a closed circle on 

the cladogram. 

Discussion of the Cladogram and Resulting 

Classification 

There are three areas on the cladogram that 

are very well defined by synapomorphies (Figure 

1, Areas A, E, and F). One area delimits the 

monandrous orchids (Area A) with 10 synapomorphies. The recognition of the monandrous 

orchids as a unit, although usually an unnamed 

one (except at the family level), is not new as 

almost all orchidologists discuss this division. 

Few, if any, of the classifications of orchids have 

dealt with the differences in such an explicit 

manner and there are some surprises such as the 

presence of elastoviscin (Table 2, t.s. 37), anther 

base insertion (Table 2, t.s. 17), and staminode 

development (Table 2, t.s. 24). 

Area E is defined by five (sometimes six, see 

Table 2, t.s. 24-9) synapomorphies and separates 

all of the Epidendroideae except Gastrodieae and 

Triphoreae. 

The third well-defined area, Area F, is somewhat of a surprise; although the Orchidoideae is 

a well known subfamily, there has been some 

discussion as to what does and does not belong 

in it. Because of the eight synapomorphies dealing with what are the most distinctive column 

and pollen characters in the family, it is easy to 

decide what should be considered part of this 

group. New characters include the presence of 

auricles (Table 2, t.s. 25). 

Five other well-defined areas are apparent. 

Area D is defined by three synapomorphies and 

delimits the subfamily Epidendroideae. This is a 

well-established taxonomic group recognized by 

all workers in the family. All three of the characters, however, are new for the group while the 

traditional characters (hard, waxy pollen aggregation and incumbent anther) are apomorphies 

found in more restricted groups within the Epidendroideae. 

Other areas of the cladogram that are welldefined are the subfamily Cypripedioideae with 

three synapomorphies, the “vandoid orchids” 

(Maxillarieae, Coelogyneae, and Vandeae of the 

Epidendroideae) and the Spiranthoideae and 

Apostasioideae with three synapomorphies each. 

There are also areas that are less well-defined. 

The cladogram contains six polytomies, several 

weak dichotomies and one subfamily that is delimited only by characters that appear more than 

once on the cladogram. For instance, although 
46 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PRA DIU Li8 Lim OEO PTE THE 

88. 

FIGURES 88, 89.-Cladograms showing two possible groupings of the Listerinae (only the 

characters important to this grouping question are illustrated-see Figure 1 and Table 3): 88, 

Listerinae is grouped with the Neottioideae (18 steps); 89, Listerinae is grouped with the 

Spiranthoideae (19 steps). 

the subfamily Epidendroideae is separated from 

the rest of the orchids easily (Area D), there are 

three polytomies and one tribe (Dendrobieae) 

that are not indicated to be monophyletic. This 

is a reflection of the poor understanding of the 

diversity of column characters within this 

subfamily. 

The Spiranthoideae is more or less well-defined and the tribes Prasophylleae and Diurideae 

are strongly characterized, but the grouping 

within this subfamily is weak. There are three 

possible groupings of the subtribes (Figures 85- 

87); two of them are equally parsimonious (Figures 85, 86) and the third is only one step longer 

(Figure 87). Of the two critical characters (35-3 

and 24-2), one (35-3) varies a great deal in the 

family (Table 4) and cannot be reliably used to 

group taxa. If we choose to group using character 24-2 then there are two equally parsimonious 

alternatives (Figures 85, 86) with the only difference being whether sectile pollen aggregation 

(35-3) developed twice within the subfamily (Fig- 
NUMBER 61 

CRA 

47 

80. 

ure 85) or developed only once and then was lost 

(Figure 86). We think that at this time there is 

no reason to have a preference between these 

two cladograms. In addition, we have no indication that the Cranichis group (Cranichideae 

sensu Dressler) is monophyletic. Although both 

Goodyerinae I and Tropidinae have character 

28-3 we have not grouped them together on the 

cladogram because this character is not believed 

to be homologous in the two groups; it is associated with different types of stipes. Because of the 

instability of the groupings within the Cranichideae we have refrained from making any nomenclatorial changes except for combining Dressler’s 

Cranichideae and Erythrodeae under the name 

of the former. 

Rasmussen (1 982:8 1) suggested placing the 

Listerinae in the Cranichideae because they both 

have fascicled roots. We examined the possibility 

of such a relationship. Figures 88 and 89 show 

that based on our characters, such an arrangement would be less parsimonious. 

Are the Apostasioideae more closely related to 

the Cypripedioideae than to the Neuwiedioideae? The two alternative arrangements are 

shown in Figures 90 and 91 and they indicate 

that both interpretations are equally parsimonious. The different arrangements depend on 
48 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

NEU APO CYP NEU APO CYP 

FIGURES 90, 91 .--Ckidograiiis showing two possible groupings of the Apostasioideae (only the 

characters important to this grouping question are illustrated-see Figure 1 and Table 3): 90, 

Apostasiodieae is more closely related to Cypripedioideae than to Neuwiedioideae (1 1 steps); 

9 1, Apostasiodieae is more closely related to Neuwiedioideae than to Cypripedioideae (1 1 

steps). 

7 

02. 32-2 38/7-31 I 

orrc 

n 

I:-3 

26-2/ 03. 

FIGURES 92-94.-Cladogranis stio\ving three possible groupings of the Neottioideae (only the 

characters important to this grouping question are illustrated; see Figure 1 and Table 3): 92, 

Xeottioideae is the sister group of the Orchidoideae/Epidendroideae (20 steps); 93, Neottioideae is the sister group of the Spiranthoideae (20 steps); 94, Neottioideae is the sister group of 

the Orchidoideae (22 steps). 
NUMBER 61 49 

whether one wishes to postulate a parallel origin 

for the development of the outer median staminode (Table 2, t.s. 16) or for the operculate 

colpus (Table 2, t.s. 42). We have chosen to 

illustrate the Apostasioideae as more closely related to the Cypripedioideae than either is to the 

Neuwiedioideae because we have more confidence in the staminode character (Table 2, t.s. 

16). We have not been able to find the operculate 

colpus (Table 2, t.s. 42) in Apostasia even though 

Schill (1 978) reports that it exists. The Cypripedioideae appear to have an annulus which may 

represent a reduced operculum. Regardless of 

which pattern one prefers, it is obvious that the 

Apostasioideae are not as closely related to the 

genus Neuwiedia as has been indicated in past 

classifications where they were placed in the same 

subfamily. 

Without a doubt, the most interesting area on 

the cladogram is the placement of the Neottioideae. There are three nearly equally parsimonious distributions of apomorphies. In one, the 

I II 

38 t 

32-24 7-34 

94. 

Neottioideae is positioned on the cladogram so 

that it is the sister group of the Orchidoideae/ 

Epidendroideae (Figure 92). It can also be drawn 

as the sister group of the Spiranthoideae (Figure 

93) with which it shares cohesion strands (character 38), and finally as the sister group of the 

Orchidoideae (Figure 94) with which it shares a 

basal viscidium attachment (character 7-3). The 

last one is two steps longer than the other two 

and is not considered further. Of the first two 

we have selected the first to use in our main 

diagram because it does not cause parallelisms in 

what appears to be a very stable character (Table 

2, t.s. 17). However, the Neottioideae are still a 

definable group no matter which of the two most 

parsimonious cladograms one selects (Figures 92, 

93). The possibility that they can be grouped in 

one of two ways merely emphasizes the need to 

recognize the Neottioideae as a separate subfamily. All of the defining characters of the Neottioideae are shared with either the Orchidoideae 

or the Spiranthoideae, or some section thereof, 

and one might be tempted to speculate on a 

hybrid origin for the Neottioideae involving the 

Orchidoideae and Spiranthoideae. However, hybridization occurs on the individual level and not 

on the subfamily level, so any hypothesis of hybridization would have to involve individuals and 

therefore cannot be addressed in this treatment. 

Should some groups within the Orchidaceae 

be recognized as separate families? For instance, 

Vermeulen ( 1966) recognized three families, 

Apostasiaceae, Cypripediaceae and Orchidaceae 

(monandrous orchids only), and Schlechter 

(1 9’70-1 984), two families, Apostasiaceae (including the Cypripedioideae) and Orchidaceae 

(monandrous orchids). Based on the cladogram 

there are three choices: all within one family; 

four families (Neuwiediaceae, Apostasiaceae, Cypripediaceae, and Orchidaceae-monandrous 

orchids); or three families (Neuwiediaceae, Apostasiaceae (d.iandrous orchids), and Orchidaceae 

(monandrous orchids). All of the groups are 

monophyletic and all are acceptable to most cladists. The goal of cladistic classification, however, 

is to achieve the maximum amount of informa- 
50 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

tion with the minimum amount of change. This 

goal can be rephrased as follows: maintain 

monophyletic groups while changing the existing 

nomenclature as little as possible. The second 

part of the goal applies to this situation. In order 

to justify a change in the current classification 

(and certainly one would have to be made because Neuwiedioideae would have to be recognized separately) the group or groups in question 

must be more closely related to something else 

than they are to the rest of the orchids. The 

three synapomorphies that unite all orchids indicate that all of the orchids are more closely 

related to one another than they are to anything 

else; therefore there is no information to be 

gained by breaking them up into separate families. It is our opinion that they should be maintained as a familv. 

Keys to the Subfamilies and Tribes of the Orchidaceae 

Key to the Subfamilies 

1. Anthers 2 or 3, base situated below stigma base; filaments and style fused 

only at base or filaments fused to style; pollen in monocolpate monads; 

stigma 3-lobed, situated between or above the anthers; labellum slightly 

zygomorphic or well developed. .............................. .2 

2. Anthers 3, elongate; filaments and style fused only at base; ......... 

......................................... NEUWIEDIOIDEAE 

3. Staminode, if present, filament-like; stigmatic surface radial to 

...................................... APOSTASIOIDEAE 

3. Staminode large, well developed; stigmatic surface abaxial; labellum 

well developed .......................... CYPRIPEDIOIDEAE 

1. Anther 1, base situated at or near stigma base or at, near, or above stigma 

apex; filaments, style and stigma fused into a column; pollen mostly in 

monoporate tetrads, aggregated into pollinia; stigma 2- (rarely 3-) lobed, 

abaxial, third lobe modified into a rostellum; labellum usually well developed .................................................... 4 

4. Anther parallel to stigma and rostellum; base positioned at or near 

stigma base; rostellum usually well developed and usually longer than 

broad (or reduced) .......................... SPIRANTHOIDEAE 

4. Anther extending above stigma; base at or near or above stigma apex; 

rostellum as long as broad and curled over or strap-like or reduced 

(rarely absent). ......................................... .5 

5. Anther base at or near stigma apex; rostellum reduced to absent or 

sensitive on contact, rarely longer than broad (Listerinae); pollinia 

granulate, in monads or tetrads ............... NEOTTIOIDEAE 

5. Anther base above stigma; rostellum strap-like or curled over; pollinia sectile or hard and waxy (rarely granulate), in tetrads (rarely 

monads) ............................................. 6 

6. Column erect or bending backwards; anther erect or at right 

angles to column or upside down, fused to column apex, locules 

2. Anthers 2, short and broad; filaments fused to style; ............ .3 

slightly asymmetric; labellum slightly zygomorphic ............. 
NUMBER 61 51 

separated by a connective; pollinia sectile with elastoviscin caudicles ................................ ORCHIDOIDEAE 

6. Column erect or bending forward; anther incumbent and free 

(rarely suberect), locules covered by cap-like connective; pollinia 

usually hard and waxy (rarely sectile or granulate); often with 

accessory structures .................... EPIDENDROIDEAE 

Keys to the Tribes 

SPIRANTHOIDEAE 

1. Filament and style greatly reduced, stigma and anther free; rostellum 

highly reduced; staminodes free, long, thin; pollinium granulate ....... 

................................................ DIURIDEAE 

1. Filament, style and stigma fused and not highly reduced; rostellum longer 

than wide; staminodes free or fused to sides of column; pollinia usually 

sectile ................................................... 2 

2. Staminodes free, long, thick; caudicle reduced, sterile: stipe hamular; 

clinandriuni incompletely fused ................ PRASOPHYLLEAE 

2. Staminodes fused to column and inconspicuous; caudicle reduced, sterile; stipe absent (except in Tropidinae); clinandrium completely fused 

........................................... CRANICHIDEAE 

NEOTTIOIDEAE 

1. Staminodes inconspicuous or absent; rostellum reduced, absent or sensitive ............................................ NEOTTIEAE 

1. Staminodes well-developed; rostellum reduced and usually removed with 

viscidium ................................................ .2 

2. Column extremely reduced or absent; staminodes fused into a high 

collar-like structure: filament long and fused to inside of staminodal 

hood ...................................... THELYMITREAE 

2. Column long and ~.ell-developed; staminodes well-developed, broad 

and fused to sides of column; filament reduced ................ .3 

3. Staminodes flared (resembling hood of cobra); stigma at apex of 

column .................................. GEOBLASTEAE 

3. Staminodes wing-like; stigma midway down column ............ 

...................................... PTEROSTYLIDEAE 

ORCHIDOIDEAE 

1. Column erect: anther erect: stigma concave ............. ORCHIDEAE 

1. Column bending backward; anther upside down or at right angles to 

column; stigma not concave. ................................. .2 
52 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

2. Anther upside down and below stigma; column turned over at apex 

.............................................. SATYRIEAE 

2. Anther at right angles to stigma;* column bent over only at anther 

................................................. DISEAE 

EPIDENDROIDEAE 

1. Pollinia sectile ............................................ .2 

2. Anther suberect .............................. GASTRODIEAE 

2. Anther incumbent ............................. ARETHUSEAE 

1. Pollinia granulate or hard and waxy ........................... .3 

3. Anther suberect ............................... TRIPHOREAE 

3. Anther incumbent ...................................... .4 

4. Pollinia granulate ............................ VANILLIEAE 

(also Hexalectris and Sobralia in the Epidendreae) 

4. Pollinia hard and waxy ................................ .?I 

5. Caudicles as many as pollinia, embedded with pollen; viscidium 

semi-liquid or rarely solid ................... EPIDENDREAE 

5. Caudicles usually sterile or absent, if pollen embedded highly 

reduced or fused; viscidium absent or solid .............. .6 

6. Pollinia with attenuate apex; viscidium solid ............. 

Pleurothallis group. ................................ 

6. Pollinia rounded, rarely attenuate, naked or with stipe; viscidium absent or solid ............................. .7 

7. Pollinia naked, rarely with a hamular stipe (Bulbophyllum), 

andsideby side ............................... 8 

8. Pollinia in sets, equal .............. DENDROBIEAE 

8. Pollinia is sets, unequal ............. MALAXIDEAE 

7. Pollinia with tegular stipe and sterile caudicle or fused 

pollen embedded caudicles with or without stipe, superposed ....................................... 9 

9. Caudicles fused, pollen embedded; with or without 

9. Caudicles free, sterile, reduced; with tegular stipe . . 10 

10. Pollinia free in sets .............. MAXILLARIEAE 

10. Pollinia fused in sets .................. VANDEAE 

tegular stipe ..................... COELOGYNEAE 

* Dressler (pei-s. coiiitii., 1983) notes that "Dim does not al\cays seem to have an anther at right 

angles to the stigina." 
NUMBER 61 

Classification of the Orchidaceae 

Non-Monandrous Orchids 

NEUWIEDIOIDEAE Burns-Balogh and Funk, 

new subfamily 

PLATE la 

Anthereae tres, abaxiales, insertae infra stigma. Filamenta et stylus connati basi. Pollinis 

grana operculata. 

Anthers 3, abaxial, representing outer median 

stamen and inner lateral stamens, free, elongate, 

base below stigma base; filaments long, fused at 

base to style. Pollen in monads, semi-tectate, with 

tectum, with columellae, foot layer present, operculate, monocolpate. Stigma lobes three, radially arranged, style equal to or longer than filaments. Perianth tubular, rudimentary nectar sac, 

median petal slightly irregular. Fruit a capsule or 

berry, 3-chambered, axile placentation. 

TypE.--Neuwiedia Blume in Hoeven and 

Vriese, Tijdschr. nat. geschied. 1. 1834, p. 142. 

DISTRIBuTI0N.-Malaysia. 

TAXA.-~ genus with 10 species. 

APOSTASIOIDEAE 

PLATE 1 b 

Anthers 2, abaxial representing inner lateral 

stamens, broad, long, adherent at margins forming a tube around style, base below stigma base; 

filaments short, fused to style; staminode, when 

present, filament-like, adherent to anther margins. Pollen in monads, semi-tectate, with tectum, 

with columellae, foot layer present, operculate, 

monocolpate. Stigma 3-lobed, radially arranged, 

exserted; style longer than anthers. Perianth radiating, nectar sac absent, parts more or less 

similar. Fruit an elongate capsule, 3-chambered, 

axile placentation. 

DISTRIBuTroN.-Tropica~ Asia, Malaysia, Australia. 

TAXA.-~ genus with ten species. 

CYPRIPEDIOIDEAE 

53 

PLATES lc-e, 3a 

Anthers 2, abaxial, representing inner lateral 

stamens, separate at each side of column, 

rounded, base below stigma base; filaments short, 

fused to style; staminode large, well-developed; 

pollen in monads, mass sticky or powdery, tectate-imperforate, with incipient columellae, foot 

layer absent, monocolpate. Stigma 3-lobed, abaxial, facing towards labellum, longer than anthers, 

style thick. Perianth zygomorphic with well-developed labellum and nectary, outer lateral sepals 

often fused to form a flag-like structure above 

labellum. Fruit an elongate capsule, 3-chambered (Selenipedium, Phragmipedium) or 1 -chambered (Cjpripedium, Paphiopedilum), placentation 

axile or parietal. 

pripedium: northern temperate, 50 species; 

Phragmipedium, tropical South America, 11 species; Selenipedium, tropical South America and 

West Indies, 3 species; Paphiopedilum, tropical 

Asia, 50 species. 

DISTRIBUTION AND TAxA.-Four genera: cyMonandrous Orchids 

Anthers 1, abaxial, representing median outer 

stamen, free or fused, elongate, parallel to or 

above stigma; filament with some portion of apex 

free or fused; staminodes 2, representing inner 

lateral stamens. Pollen in tetrads, rarely monads, 

united into masses called pollinia, exine various, 

pore one, proximal; with elastoviscin. Stigma 

fused to style and filaments to form a central 

column structure, lobes 2, some with a portion 

of third lobe also receptive; rostellum representing modified third stigma lobe (apical); viscidium 

present in most genera. Perianth zygomorphic, 

nectary usually present, labellum well-developed 

in most genera. Fruit an elongate capsule, ovary 

1 -chambered, placentation axile. 

This study is only to the tribal level, though in 
54 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

some cases we have been able to define the 

subtribes also. For the most part our characters 

are apomorphic on the tribal level. Because of 

this most of the subtribes listed here are those of 

Dressler (1 98 1). 

SPIRANTHOIDEAE 

Anther base situated at or near base of stigma, 

rarely near apex, free, erect to suberect, dorsal, 

parallel to stigmatic surface and rostellum; clinandrium well-developed, composed of the back 

of the stigma and margins of staminodes; staminodes inconspicuous and fused with column apex 

to well developed and free. Pollen in granulate 

or sectile masses, tetrads, cohesion strands, semitectate to tectate, foot layer, columellae, and 

tectuni present. Stigma 2-lobed; rostellum erect 

to suberect, apical portion abscissing and forming 

the viscidiuni; viscidium mostly apical to rarely 

subapical depending on rostellum development; 

column foot rarely well developed. (See “Monandrous Orchids” for additional characters.) 

DIURIDEAE 

PLATE If 

Staminodia membranous, free, prominent; clinandrium incompletely fused; pollinarium granulate, composed of monads (less often loosely 

aggregated tetrads); column absent or extremely 

reduced; stigma free, not fused to column, rostellum removed with viscidium portion; rootstem tuberoids. 

DIsTRrBUTIoN.-Austra~ia. 

TAXA.-2 genera (Diuris, Orthoceras); 40 species. 

PRASOPHYLLEAE (Schlechter) Burns-Balogh and 

Funk, new status 

PLATE lg 

Type: Prasophyllum R. Brown. Prodromus 

Florae Novae Hollandiae. 18 10. p.3 17.-Staminodes free, long, thick; clinandrium incompletely 

fused; pollinia sectile, rarely granular, hamular 

stipe present, with reduced sterile caudicles; rostellum well-developed and longer than broad; 

flowers non-resupinate. 

TypE.-Prasophyllum R. Brown, Prodromus 

Florae Novae Hollandiae. 18 10, p. 3 17. 

DIsTRIBuTIoN.-Austra~ia, New Zealand. 

TAXA.--~ genus with 40 species. 

CRANICHIDEAE 

PLATES lh-k. 3b-i 

Staminodes short and inconspicuous; clinandrium completely fused; pollinia sectile or granulate, with or without accessory structures; rostellum well-developed, longer than broad; flowers resupinate or non-resupinate. 

TAXA.-7 subtribes. 

GOODYERINAE 

Tegular stipe and reduced sterile caudicle in 

some species; stems succulent trailing, rooting at 

the nodes. Three species of Goodyera have an 

incumbent anther above stigma. 

DISTRIBUTION.-Most~y tropical but also temperate. 

TAXA.-~~ genera; 425 species. 

TROPHIDINAE 

Hamular stipe and reduced sterile caudicle 

present; sectile, rarely secondarily granulate pollinia; stems reed-like, leaves plicate. 

DIsTRIBUTION.-Pan tropical. 

TAXA.-2 genera; 12 species. 

SPIR ANTHIN AE-I 

Sterile caudicles rarely present; pollinia in unDIsTRrBuTIoN.-Mostly tropical America. 

TAXA.--~ 7 genera; 325 species. 

equal sets. 

MANNIELLINAE-I I I 

Staminodes prominent. 

DISTRIBUTION.-Africa. 

TAXA.--~ monotypic genus. 
NUMBER 61 

CRANICHIDINAE-I1 

Flowers non-resupinate; some with hamular 

DIsTRIBUTION.-Tropical and temperate 

TAXA.-~ 5 genera; 200 species. 

stipe; pollinia round in cross section. 

areas. 

55 

LISTERINAE 

CRYPTOSTYLIDINAE-I 

Pollinia in unequal sets. 

DrsTRIBuTroN.-Austra~ia and Asia. 

TAXA.- 1 genus with 15 species. 

PACHYPLECTRONINAE-I11 

Staminodia web-like between stigma and anDISTRIBUTION.-New Caledonia. 

TAXA.--I genus with 2 species. 

ther margins. 

Pollen grain pore distal or equatorial; rostellum sensitive, viscidium attachment above base 

of pollinia but rarely at apex; anther base in 

Neottia varies from near base of stigma to apex 

of stigma. 

DISTRIBUTION.-North temperate. 

TAXA.-2 genera; 38 species. 

LIMODORINAE 

Rostellum absent or reduced, when present 

viscidium basal or just above base; anther base 

varies in Epipactis from near base of stigma to 

apex of stigma. 

DIsTRIBuTIoN.-Austra~ia, tropical Africa and 

Asia, and northern hemisphere. 

TAXA.-5 genera including Rhizanthella and 

Cryptanthemis; 62 species. 

NEOTTIOIDEAE 

THELYMITREAE 

Anther base inserted at or near apex of stigma, 

free, erect to suberect or nodding, free, elongate; 

filament often bending over so anther appears 

terminal, free or fused to staminodes; pollen in 

granulate masses, monads or tetrads, with cohesion strands, semi-tectate to tectate, tectum, foot 

layer, and columellae present, with proximal or 

equatorial or distal pore. Stigma 2-lobed, usually 

concave in those with long column; rostellum 

short or extremely reduced or absent or sensitive; viscidium short and round or absent or semisolid; column foot not well-developed. (For additional characters, see section on “Monandrous 

Orchids,” p. 53.) 

TAXA.-4 tribes. 

NEOTTIEAE 

PLATE 3j 

Staminodes reduced or absent; pollinium soft 

and mealy; rostellum, if present, greatly reduced 

or sensitive on contact. 

TAXA.-2 subtribes. 

PLATE lm 

Staminodes fused into a high collar-like structure and fused to the filament; pollinium soft 

and mealy; column highly reduced, rostellum 

very short, viscidium solid. 

DIsTRIBUTIoN.-~a~aysia, Australia, and New 

Zealand. 

TAXA.-~ genera (Thelymitra, Calochilus, and 

Epiblema); 60 species. 

GEOBLASTEAE 

PLATE 11 

Staminodes broad and fused to sides of column 

forming a cobra-hood-like structure below and 

partially at base of anther and tapering to base 

of column, resembling some Epidendroideae; 

pollinia soft and mealy, rarely divided (Caladenia); column long and well-developed; rostellum 

reduced and usually removed with viscidium portion, viscidium solid to semi-solid. 

TAXA.-2 subtribes. 
56 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

CHLORAEINAE 

DISTRIBUTION.-south America and New CaTAXA.-6 genera; 100 species. 

ledonia. 

CALADENIINAE 

DIsTRIBuTI0N.-Australasian. 

TAXA.- 16 genera, 100 species. 

PTEROSTYLIDEAE (Schlechter) Burns-Balogh and 

Funk, new status 

Staminodes broad at apex and wing-like, tapering at base of column; pollinia soft and mealy; 

column long and well-developed; stigma separated from rostellum by long narrow sterile area, 

stigma oblong, positioned at 9 to % length of 

column; rostellum reduced and often removed 

with viscidium, viscidium semi-solid. 

TYPE.-Pterost$is R. Brown, Prodromus 

Florae Novae Hollandiae. 1810, p. 326. 

DIsTRIBuTI0N.-Australasian. 

TAXA.-~ genus with 70 species. 

ANOMALOUS 

Genoplesium, Microtis (hamular stipe); Drakea, 

Spiculaea (Drakaeinae!). 

ORCHIDOIDEAE 

Anther base inserted above stigma apex, fused 

to column apex, erect, inclined away from column axis, or bent backwards and upside-down, 

locules separated by anther connective; filament 

absent; clinandrium absent; staminodes with callus-like auricles. Pollen in tetrads united into 

massulae which are united by elastoviscin core 

threads; interlocular elastoviscin caudicles present on each pollinium set; pollen exine semitectate to intectate, foot layer absent, monoporate, proximal pore. Stigma 2-3 lobed, some on 

stigmaphores (Habenariinae), concave or protruding; rostellum strap-like at base, between 

anthers, often forming a bursicle; viscidium 1-2 

parted; column foot absent. (For additional characterics see “Monandrous Orchids.”) 

TAXA.-3 tribes. 

ORCHIDEAE 

PLATES In-p, 2a, 3k 

Anther erect; labellum often spurred. 

TAXA.--3 subtribes. 

ORCHIDINAE 

D1sTRIBuT1oN .-Most 1 y northern. 

T~xA.-36 genera; 600 species. 

H ABENARIINAE 

DISTRIBUTION.-Pantropical, Eurasia and AfTAXA.-2 1 genera; 1 100 species. 

rica. 

HUTTONAEINAE 

DIsTRIBuTI0N.-Southern Africa. 

TAXA.-~ genus with 5 species. 

SATYRIEAE 

Anther upside down, bent backwards and below stigma; labellum double spurred. 

TAXA.-3 genera; 1 10 species. 

D~STR~B~TION. -Mostly Africa. 

DISEAE 

PLATE 2b 

Anther at right angles to column, bending 

TAXA.--:! subtribes. 

backwards; labellum not spurred; sepaline spur. 

DISINAE 

DIsTRIBuTIoN.-Africa, especially southern 

TAXA.-~ genera; 190 species. 

Africa. 
NUMBER 61 

CORYCIINAE 

57 

STEREOSANDRINAE 

DISTRIBuTIoN.-Mostly Africa, but extending 

into tropical Asia. 

EPIDENDROIDEAE 

Anther situated above stigmatic surface, free, 

incumbent, rarely erect to suberect, with cap-like 

connective covering locules, locules usually 4 or 

divided into partitions; filament elastic or free 

and membranous or thick, or fused to column 

apex; clinandrium composed of dorsal (back) surface of rostellum and apex of column (with staminodes). Staminodes fused to column apex, some 

with free projections. Pollen in soft, mealy masses 

or hard, waxy spheres with or without accessory 

structures; exine semi-tectate to tectate, with or 

without foot layer, monoporate, with proximal 

pore. Stigma concave, tucked under rostellum, 

rarely protruding; rostellum broad and curled 

over, rarely erect, remnant often long and narrow; viscidium semi-liquid or solid or absent; 

column foot often well-developed. (For additional characters, see “Monandrous Orchids.”) 

TAXA.- 10 tribes, 1 informal group (Pleurot ha 1 1 is) . 

GASTRODIEAE 

Anther suberect; pollinia 4 in 2 sets, sectile, 

rarely secondarily granulate; pollen grain with or 

without foot layer to columellae, exine psilate to 

reticulate and no accessory structures (except 

Epipogoninae). 

TAXA.-~ subtribes. 

EPIPOGONINAE 

With stipe and solid viscidium. 

DIsTR1BuTIoN.-Eurasia, tropical Africa, tropTAXA.-~ genus with 2-3 species. 

ical Asia. 

GASTRODIINAE 

DIsTRIBUTI0N.-Pantropical, mostly tropical 

TAXA.-~ genera; 50 species. 

Asia and Australasia. 

DIsTRIBUTI0N.-Eurasia. 

TAXA.--~ genus with 2-3 species. 

W ULLSCHLAEGELIINAE 

DISTRIBUTION.-Tropical America. 

TAXA.-~ genus with 2 species. 

NERVILIINAE 

DIsTR1BuTIoN.-Tropical Asia and Africa. 

TAXA.-~ genus with 80 species. 

TRIPHOREAE 

Anther suberect, connective cap not well developed; pollinia 4, granulate, pollen grain exine 

reticulate, with or without foot layer, columellae 

absent; viscidium semi-liquid. 

TAXA.-2 subtribes. 

ACIANTHINAE 

DIsTRIBuTIoN.-AUstra~asia. 

TAXA.-4 genera; (?) 100 species. 

TRIPHORINAE 

DIsTRIBuTIoN.-TropiCa~ America to North 

TAxA.-~ genera; 20 species. 

America. 

ARETHUSEAE 

Anther incumbent; pollinia number (?), sectile; 

pollen grain exine psilate to coarsely reticulate, 

with or without foot layer; viscidium semi-liquid 

(?I. 

DISTRIBUTION.-North America. 

TAXA.- 1 monotypic genus. 

VANILLIEAE 

Anther incumbent; pollinia granulate, in monads or loosely aggregate tetrads, foot layer present; stigma with two flap-like lobes, viscidium 

semi-liquid. 

TAXA.-4 subtribes. 
58 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

VANILLINAE 

Three chambered ovary in Eriaxis and Vanilla. 

TAXA.-~ genera; 165 species. 

DISTRIBUTION.-Pantropics. 

PALM o R c H I D I N A E 

DISTRIBUTION.-Tropical America and AfTAXA.-~ genera; 12 species. 

rica. 

POCONIINAE 

DISTRIBUTION.-Tropical and North AmerTAXA.-5 genera; 40 species. 

ica. 

LECANORCHIDINAE 

Pollen polyporate. 

DISTRIBUTION.-Tropical Asia. 

TAXA.-~ genus with 20 species. 

EPIDENDREAE 

PLATES 2c-e, 31-m 

Anther incumbent; pollinia 4-8 with pollen 

embedded caudicles, pollinia side by side, 

rounded, more or less equal in size; viscidium 

semi-liquid to solid, foot layer absent. 

TAXA.-~ subtribes. 

BLETIINAE 

DISTRIBUTION.-MOStly pantropical. 

T~xA.-26 genera; 380 species. 

SOBRALINAE 

DIsTRIBuTIoN.-TropiCa~ America. 

TAXA.-5 genera; 150 species. 

LAELIINAE 

DIsTRIBUTION.-Tropical America. 

TAXA.-~~ genera; 830 species. 

PLEUROTHALLIS Group 

PLATES 2Jg, 3n, 4a 

Anther incumbent; pollinia 2-8, with narrow 

attenuate apex, viscidium solid, foot layer absent. 

TAXA.-~ subtribes. 

PLEUROTHALLIDINAE 

DISTRIBUTION.-Tropical America. 

TAXA.-26 genera; 3800 species. 

MEIRACY LLIINAE 

DISTRIBUTION.-Tropical Mexico and Central 

TAXA.-~ genus with 2 species. 

America. 

PODOCHILINAE 

DISTRIBUTION.-Australasia, tropical Asia and 

TAXA.-7 genera; 230 species. 

Africa. 

THELASIINAE 

DISTRIBUTION.-Tropical Asia and Australia. 

TAXA.-7 genera; 270 species. 

ERIINAE 

DISTRIBUTION.-Tropical Asia, Australia, and 

TAXA.-8 genera; 500 species. 

Africa. 

ADRORHIZIN AE 

DISTRIBUTION.-Tropical Asia. 

TAXA.-2 genera; 3 species. 

DENDROBIEAE 

Anther incumbent; pollinia 4, naked, rarely 

with reduced pollen caudicles, waxy, pollinia side 

by side, adherent in sets, in more or less equal 

member sets, foot layer absent; elastoviscin between pollinia in sets; viscidium absent. 

TAxA.-~ subtribes. 
NUMBER 61 5 9 

THUNIINAE 

Pollinia eight. 

DIsTRIBUTIoN.-TropiCa~ Asia. 

TAXA.-~ genera; 42 species. 

DENDROBIINAE 

DISTRIBUTION.-Tropical Asia. 

TAXA.-~ genera; 1650 species. 

GLOMERINAE 

Pollen embedded caudicles in some. 

DIsTRIBuTIoN.-TropiCa~ Asia and Australia. 

TAXA.-~ genera; 130 species. 

SUNIPIINAE 

Pollen embedded caudicles in some. 

DIsTRIBuTIoN.-TropiCa~ Asia. 

TAXA.-~ genera; 25 species. 

BULBOPHYLLINAE 

One species of Bulbophyllum with hamular 

stipe and reduced sterile caudicles. 

TAXA.-~ genera; 1020 species. 

DISTRIBUTI~N. - Pan tropics. 

MALAXIDEAE 

Anther incumbent; pollinia 4, side by side, 

naked, in unequal sets, foot layer absent; viscidi u~ii semi-liq uid . 

 DISTRIBUTION.-^ genera; 1890 species. 

CORALLORHIZINAE 

DISTRIBUTION.-North temperate and tropiTAXA.-~ genera; 60 species. 

cal America. 

MAXILLARIEAE 

Anther incumbent; pollinia 4 in unequal sets, 

superposed, elastoviscin between pollinia, caudicle sterile, tegular stipe; viscidium solid. 

TAXA.- 10 subtribes. 

CALYPSOINAE 

DISTRIBUTION.-North America and eastern 

TAXA.-2 genera; 3 species. 

Asia. 

CRYPTARRHENINAE 

Stipe absent. 

DISTRIBUTION. -Tropical America. 

TAXA.--~ genus with 3-4 species. 

ZYGOPETALINAE 

DIsTRIBuTION.-Tropica~ America. 

T~xA.-26 genera; 150 species. 

BIFRENARIINAE 

DIsTRIBuTIoN.-Tropica~ America. 

TAXA.-5 genera; 50 species. 

LYCASTINAE 

COELOCY NEAE 

Anther incumbent; pollinia 4 in equal sets, 

superposed, Fvith fused pollen caudicle and/or 

tegular stipe, foot layer absent; viscidium solid 

or semi-liquid. 

TAXA.-2 subtribes. 

DI~TRIBuTI~N, -Tropical America. 

TAXA.-~ genera; 40 species. 

MAXILLARIINAE 

DIsTRIBuTIoN.-TropiCa~ America. 

TAXA.-~ genera; 485 species. 

COELOGYNINAE 

DICHAEINAE 

Stipe absent. 

DIsTRIBUTIoN.-Tropical Asia. 

TAXA.-4 genera; 440 species. 

DIsTRIBuTI0N.-Tropical America. 

TAXA.-~ genus with 45 species. 
60 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TELIPOGONINAE 

DIsTRIBuTIoN.-Tropica~ America. 

TAXA.-4 genera; 60 species. 

ORNITHOCEPHALINAE 

DIsTRIBuTIoN.-Tropica~ America. 

TAXA.-14 genera; 70 species. 

CYRTOPODIINAE 

DISTRIBuTIoN.-Pantropical. 

TAXA.-24 genera; 425 species. 

GENYORCHIDINAE 

DIsTRIBuTIoN.-TropiCa~ Africa. 

TAXA.--~ genus with 6 species. 

POLY STACHY INAE THECOSTELINAE 

DISTRIBUTION.-PantrOpiCS, mostly Africa. DISTRIBUTION.-Tropical Asia. 

TAXA.-4 genera; 220 species. TAXA.-~ genus with 5 species. 

VANDEAE 

PLATES 2h-n, 46-k 

Anther incumbent; pollinia 4 in unequal sets 

superposed, fused (rarely free), foot layer absent, 

caudicle sterile, stipe tegular; viscidium solid. 

TAXA.-~ 1 subtribes. 

SARCANTHINAE 

DIsTRIBuTI0N.-Tropical Asia, with a few speTAXA.--~~ genera; 1000 species. 

cies in Africa. 

C ATASETINAE 

Unisexual; pollinaria spring-ejected. 

DIsTRIBuTI0N.-Tropical America. 

TAXA.-~ genera; 145 species. 

ACRIOPSIDINAE 

DIsTRIBuTIoN.-TropiCa~ Asia. 

TAXA.--I genus with 12 species. 

STANHOPAEINAE 

DISTRIBUTION.-Tropical America. 

TAXA.-~ 7 genera; 190 species. 

AER ANGIDINAE 

PACHYPHYLLINAE 

Double stipe in some. 

DIsTRIBuTI0N.-Tropical Africa. 

TAXA.-~~ genera; 300 species. 

ANGRAECINAE 

DISTRIBUTION.-Tropical Africa and tropical 

TAXA.-16 genera; 400 species. 

America. 

Double stipe in some. 

DIsTRIBuT1oN.-Tropical America. 

TAXA.-2 genera; 25 species. 

ONCIDIINAE 

DIsTRIBuTIoN.-TropiCa~ America. 

TAXA.-~~ genera; 950 species. 

Conclusion 

Our system of classification follows the principles of phylogenetic systematics in that the emphasis is on the recognition and maintenance of 

monophyletic sister groups. We have used only 

characters that we found to be consistent at the 

tribal or subtribal level. Further, we have used 

only apomorphies to make groups (construct the 

cladogram). The resulting hierarchy is based on 

the cladogram and all recognized groups are 

monophyletic unless otherwise indicated. 
NUMBER 61 61 

Major points of comparison with other systems 

are as follows: 

1. Our system differs from all others in the 

establishment of the subfamily Neuwiedioideae 

containing only the genus Neuwiedia. 

2. The subfamily Spiranthoideae differs from 

that of Dressler in that we have transferred, as 

tribes, the Prasophyllinae (sensu Burns-Balogh, 

1984c) and the Diurideae (sensu Burns-Balogh 

and Funk, 198413) into this subfamily from the 

Orchidoideae (by Dressler, 198 1). 

3. Our concept of the Neottioideae is somewhat different from any other system in that we 

have transferred into it most of Dressler’s Diurideae (except those referred to in no. 2 above) 

and the tribe Neottieae. We have resurrected 

Pfitzer’s Thelymitreae with a single genus, and 

have raised the rank of the subtribe Pterostylidinae to tribal level. Caladeniinae and Chloraeinae have been combined into the Geoblasteae. 

Acianthinae, treated as a subtribe of Diurideae 

by Dressler, has been tentatively transferred to 

Triphoreae (Epidendroideae). 

4. The tribes of the Epidendroideae presented 

some problems because of the lack of information on the column structure. However, several 

changes were made. The four tribes that remain 

similar to those of Dressler are Gastrodieae, Vanillieae, Malaxideae, and Dendrobieae. The Arethuseae, Triphoreae, Epidendreae, Pleurothallis 

Group, Coelogyneae, Maxillarieae, and Vandeae 

have been rearranged as far as contents of the 

subtribes. This is a tentative arrangement reflecting the information that was available for this 

study. Of all the areas on the cladogram we think 

these groupings have the greatest chance of 

changing as more information becomes available. 

The subtribes are those of Dressler. We could 

not cladistically demonstrate that four of these 

are monophyletic, however, because we have no 

synapomorphies for them (see Figure 1). 

We feel that this classification is based on consistent characters and therefore reflects the natural groups within the Orchidaceae. However, 

we view this classification as a step in the direction 

of a phylogenetic classification because there are 

areas on the cladogram that lack resolution. Certainly, there is a need for specialists in the various 

groups to work within each tribe to determine 

the naturalness of the subtribes. New data in the 

form of additional synapomorphies or reinterpretation of the transformation series used in this 

study may indicate changes in this classification, 

especially in the subfamily Epidendroideae. One 

of the most advantangeous aspects of cladistic 

methodology is that it allows for easy criticism 

and reevaluation. We hope that this classification 

will provide the stimulation for those who both 

agree and disagree with our method and results 

to provide additional information to test our 

classification. 
APPENDIX 

List of Specimens Examined 

The data in this section are presented by 

subfamilies. The species name is followed by the 

collector’s name and number. The herbarium or 

collection where the specimen is housed and the 

collection number, when available, are given in 

parentheses. Abbreviations are as follows: AM0 

= Associacion Mexicana de Orquideologia; NYS 

= New York State Museum, Albany, NY.; S1 = 

U.S. National Orchid Collection, National Museum of Natural History, Smithsonian Institution, Washington, D.C.; SEL = Marie Selby Botanical Garden Greenhouse; NCBG = National 

Captial Botanical Garden Greenhouse, Canberra; s.n. = sine numero (without number). 

These greenhouse plants often had no collector 

or collecting number. If the abbreviation is followed by a number, then it indicates the greenhouse identification number and not the collecting number. Balogh = preserved material held 

in Balogh’s personal collection. 

APosTAs1oIDEAE.-Apostasia Sp., Booea 7683 

(US); A. wallichii, Brass 25570 (US), Booea 7808 

(US). 

NEUWIEDIOIDEAE.-NeUWiedzU zollingeri, 

Booea 7949 (US); N. veratijiolia, Booea 8991 

(US); N. grijijthii, Bartlett 6421 (US), Yates 2258 

(US). 

CYPRIPEDIo1DEAE.-Cypripedium acaule, Balogh 1053 (US); C. xmontanum, Balogh 1059 (US); 

C. pubescens, Dressler s.n. (Balogh). Paphiopedilum glaucophyllum (SI 78 1862, SEL 1-75-77); 

P. exul (SI 781 138); P. maudiae, cultivated (Balogh); P. spicerianum (SI 78 108 l). Phragmipedium 

caricinum (SI 7843 1); P. longijiolium, Dressler s.n. 

(Balogh), Dodson s.n. (SEL); P. schlimi, Mejia sen. 

(Balogh). Selenipedium chica, Dressler s.n. (Balom. 

SPIRANTHo1DEAE.-Altensteiniujmbriata, Barriga 7769 (US). Anoectochilus regalis, Jayasuriya 

Baskeruillea paraensis, Handro 2256 (US). Beloglottis costaricensis, Greenwood 858 (AMO). 

Brachystele afjnis, Greenwood 865 (AMO); B. 

polyanthus, Greenwood 2 (AMO), 84 1 (AMO). 

Cheirostylis gymnochiloides, Humbert 17794 

(US); C. longilora, Smith 1628 (US). Corymborkis 

Java, Killip and Hazen 8982 (US); C. forcipigera, 

Turckheim 860 (US). Cranichis apiculata, Greenwood 793 (AMO); C. cililabia, Greenwood 783 

(AMO); C. cucullata, Greenwood 825 (AMO); C. 

subumbellata, Suarez 92 (US). Cryptostylis subuLata, Bernhardt s.n. (Balogh). Cyclopogon congestus, Regnell 1197 (US); C. elatus, Greenwood 

519 (AMO); C. millei, Luer 2414 (SEL); C. saccatus, Arsene 5152 (US); C. sp., Foster s.n. (US). 

Deiregyne hemichrea, Greenwood 752 (AMO), 

Hagsater s.n. (AMO); D. obtusa, Greenwood 32 1 

(AMO); D. pulchra, Greenwood 407 (AMO); D. 

pyramidalis, Greenwood 798 (AMO), Balogh 

1019 (US); D. tenuiflora, Greenwood s.n. (AMO), 

Dichromanthus cinnabarinus, Greenwood 990 

(AMO), Balogh 723 (US). Diuris aurea, Clements 

2384 (NCBG), D. longijiolia, Clements 2108 

(NCBG); D. maculata, Bernhardt s.n. (Balogh), 

Clements 2388 (NCBG); D. sulphurea, Clements 

2389 (NCBG). 

Erythrodes sp., Foster s.n. (Balogh); E. sp., 

Dressler 4675 (US); E. hyphaenctica, Dodson s.n. 

(SEL); E. maculata (SEL 81-1771); E. procera, 

Abbott 2705 (US). Eurystyles sp., Kuhn s.n. 

(SEL). 

Gomphicus viscosa, Wurdack 1734 (US). Goodyeru oblongijiolia, Balogh 1055 (US); G. pubescens, 

Balogh 907 (US). 

1219 (US). 

62 
NUMBER 61 63 

Hetaeria oblongijolia, Merrill 4026 (US). 

Ligeophila jamesonii, Dodson s.n. (SEL); L. stigmatoptera, Fernandez 2 14 1 (US). Ludisia discolor, 

cultivated (Balogh). 

Manniella gustaui, Tryon 6485 (AMES). Macodes anderiana (SI 781 346). 

Pachyplectron neocaledonia, McPherson s.n. 

(Balogh). Pelexia funckiana, Balogh and Greenwood 959 (US), Behib 707 (Balogh); P. laxa, 

Balogh s.n. (Balogh). Physurus sagreanus, Jimenez 

2757 (US). Platythelys sp., Dressler s.n. (Balogh). 

Ponthieua sp., Greenwood 994 (AMO); P. maculata, Greenwood 1008 (US); P. racemosa, Greenwood 1020 (AMO); P. tuerkheimii, Greenwood 

1006 (AMO). Prasophyllum sp., Clements 21 11 

(NCBG); P. alpinum, JT 1275 (NCBG); P. densum, Clements 2319 (NCBG); P. elatum, Clements (NCBG); P. jimbriatum, Clements 2301 

(NCBG); P. macrostachyum, George s.n. (Balogh); 

P. morrisii, Clements 23 16 (NCBG); P. nublingii, 

SH 1147 (Balogh); P. rufum, Clements 2318 

(NCBG); P. ruppii, Clements 2298 (NCBG); P. 

suttonii, JT 1340 (Balogh); P. woolsii, Clements 

23 12 (NCBG). Prescottia stachyoides, Dressler s.n. 

(Balogh); P. tubulosa, Greenwood 922 (AMO). 

Pterichis triloba, Ferreyra 7534 (US). 

Sarcoglottis sp., Greenwood 91 5 (AMO); S. sp., 

Greenwood 657 (AMO); S. sp., Greenwood 915 

(AMO); S. sp.,Greenwood 657 (AMO); S. sp., 

Greenwood 352 (AMO); S. sp., Greenwood 757 

(AMO); S. acaulis, Greenwood 657 (AMO); S. 

cerina, Dix 785 (Balogh); S. gutturosa, Greenwood 722 (AMO); S. rosulata, Greenwood 450 

(AMO); S. schaffneri, Hagsater 5458 (AMO), 

Greenwood 1023 (AMO). Schiedeella chloraeformis, Greenwood 317 (AMO); S. densijlora, 

Greenwood 10 1 1 (AMO); S. eriophora, Balogh 

101 1 (US), Greenwood sen. (AMO); S. falcata, 

Balogh 969 (US); S. hyemalis, Hagsater 5739 

(AMO); S. llaveana, Balogh and Greenwood 924 

(US), Greenwood 930 (AMO); S. michuacana, 

Greenwood 8 13 (AMO); S. pseudopyramidalis, 

Balogh and Greenwood 917 (US); S. saltensis, 

Greenwood 860 (AMO). Solenocentrum costaricensis, Dressler 584 1 (Balogh). Spiranthes cernua, 

Balogh 914 (US), Sheviak 1695 (NYS); S. lacera, 

Balogh 904 (US); S. ochroleuca, Balogh 913 (US), 

Sheviak 1657 (NYS); S. praecox, Grelen s.n. (Balogh); S. romanzoffiana, Pierce 91 l (Balogh). Stenoptera pilfera, Haught 33 15 (US). Stenorrhynchos 

arechavaletanii, Smith and Klein 122 17 (US); S. 

aurantiacus, Balogh 735 (US); S. balanophorostachys, Gehrt (US); S. canterae, Herter 9878 (US); 

S. cuspidata, Dodson s.n. (SEL); S. densus, Hatschbach 26598 (US); S. lanceolatus, Greenwood 393 

(AMO), Balogh 723 (US); S. lateritus, Hoehne 

and Gehrt (US); S. regius, Macedo 2667 (US); S. 

speciosus, Greenwood 5 18 (AMO); Tropidia angulosa, Henry 1905 (US). 

Vrydagzynea micronesiaca, Glassman 2860 (US). 

Zeuxine stricta, Smith 8457 (US). 

NEOTTIOIDEAE.-Aphyllorchis pallida, Clements 323 (US). Asarca sinuata, Senn 4277 (US); 

A. patagonica, Joseph 1072 (US). 

Caladenia angustata, Bernhardt s.n. (Balogh); 

C. angustijolia, Bernhardt s.n. (Balogh); C. cairnsiana, Clements 2 183 (Balogh); C. catenata, Clements s.n. (NCBG), Rogers s.n. (Balogh); C. caerulea, Bernhardt s.n. (Balogh); C. deformis, Bernhardt s.n. (Balogh); C. dilatata, Clements 2107 

(NCBG), Kelly, s.n. (Balogh) C. doutchae, Clements 2099 (NCBG); C. jilamentosa, Clements 

2102 (NCBG), Bernhardt s.n. (Balogh); C. graminfolia, Clements 2103 (NCBG); C. Java, 

Clements 2 100 (NCBG); C. latfolia, Clements 

2105 (NCBG); C. patersonii, Bernhardt sen. (Balogh); C. reticulata, Bernhardt s.n. (Balogh); Caleana minor, Matthews s.n. (US). Calochilus campestris, Clements 2386 (NCBG); C. robertsonii, 

Bernhardt s.n. (Balogh); Kelly s.n. (Balogh). Cephalanthera oregana (austinae), Sandberg and 

Leiberg 571 (US); C. rubra, Podpera 21 (US); C. 

grandzjlora, Cintract 1886 (US). Chiloglottis gunnii, Bernhardt s.n. (Balogh). Chloraea crispa, 

White s.n. (US); C. sp., Piquilieu 3745 (US); C. 

nudilabia, Hutchison 297 (US). Codonorchis lessonii, Goodall 433 (US). 

Elythranthera brunonis, Clements (NCBG). Epiblema grandzjlora, Pritzel 100 (US). Epzpactis gigantea, Hall 2370 (US), Heller and Kennedy 

887 1 (US), E. palustris, Kurzweil s.n. (Balogh), 

E. atrorubens, Stephanoff and Georgieff (US). 
64 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Eriochilus cucullatus, SH 1146 (Balogh), Bernhardt s.n. (Balogh); E. fimbriatus, French s.n. 

Genoplesium baueri, Clements 2305 (NCBG). 

Glossodia major, Mueller s.n. (US), Clements 

2348 (NCBG), Bernhardt s.n. (Balogh); G. minor, 

Boorman s.n. (US). 

Limodorum abortivum, Franchschi 343 (US). 

Listera auriculata, Fernald and Collins 200 (US); 

L. borealis, Calder and Savile 9939 (US); L. CUUTina, Parks and Tracy 11667 (US); L. ovata, 

Kurzweil s.n. (Balogh). 

Megastylis gigas, Mckee 7755 (US), Buchholz 

1186 (US). Microtis alba, Pritzel 922 (US); M. 

parvzflora, Bernhardt s.n. (Balogh); M. unzjZora, 

Fosberg 30757 (US). 

(US). 

Neottia nidus-avis, Fosberg 35805 (US). 

Pterostylis baptistii, Bernhardt s.n. (Balogh); P. 

cucullata, Clements s.n. (NCBG); P. decurva, 

Bernhardt s.n. (Balogh); P. foliata, Allison s.n. 

(US); P. nana, Bernhardt s.n. (Balogh); P. nutans, Bernhardt s.n. (Balogh); P. obtusa, Bernhardt s.n. (Balogh); P. parvzjlora, Clements 2307 

(NCBG); P. pulchella, Clements 2321 (NCBG); 

P. revoluta, SH 1160 (Balogh); P. Tufa, Clements 

s.n. (NCBG); P. truncata, Bernhardt s.n. (BalRhizanthella gardneri, George 157 17, McGuiness s.n. (US). 

Thelymitra antennijera, Bernhardt s.n. (Balogh); T. carnea, Clements 2383 (NCBG); T. media, Bernhardt s.n. (Balogh); T. nuda, Clements 

(NCBG), T. pauciflora, Bernhardt s.n. (Balogh), 

Kelly s.n. (Balogh). 

ORCHIDOIDEAE: Aceras anthrophorum, Dressler 

s.n. (Frankfurt Botanical Garden). Amerorchis rotundijolia, Sandberg 10 14 (US). Anacamptis pyramidalis, Fosberg 32759 (US), Castella 1886 

Brachycorythis pubescens, Leipoldi s.n. (US), 

Wood s.n. (US). Brownleea caerulea, Kemp 771 

Ceratandra bicolor, Schlechter 9296 (US). Chamorchis alpina, Fosberg 33093 (US), Aryet 9800 

(US), Kurzweil s.n. (Balogh). Coeloglossum viride, 

Larsen 20867 (US). Corycium nigrescens, Bayliss 

(US). Cynorchis angustipetala, Decary 61 59 (US). 

ogh). 

(US). 

(US). 

Dactylorhiza aristata, Murie 6 (US); D. incarnata, Kurzweil s.n. (Balogh). Disa bivalvata, 

Kemp 771 (US), Fourcave 529 (US); D. cornuta, 

Wright 13 1 (US); D. crassicornis, Wood 8430 

(US). Disperis sp., Ramoorthy 1920 (US), D. cooperi, Wood 8096 (US). 

Galearis spectabilis, Balogh 907 (US). Gymnadenia odoratissima, Kurzweil s.n. (Balogh). 

Habenaria alata, Greenwood 576 (US); H. clypeata, Greenwood 9 17 (AMO); H. dilatata, Balogh 1058 (US); H. distans, Greenwood 720 

(AMO); H. genuflexa, Kurzweil s.n. (Balogh); H. 

guadalajarana, Greenwood 700 (AMO); H. longiauriculata, Kurzweil s.n. (Balogh); H. macroceratitus, Greenwood s.n. (AMO); H. novemfida, 

Greenwood 89 1 (AMO); H. odontopetala, Greenwood 532 (AMO); H. oreophila, Greenwood 798 

(AMO); H. trzfda, Greenwood 719 (AMO); H. 

sp., Balogh 750 (US). Herminium monorchis, 

Kurzweil s.n. (Balogh). Herschelia homalopetala, 

Stolz 127 (US). Himantoglossum hircinum, Dvorak 

and Novacek 197 (US). Holothrix praecox, Tebrum 7850 (US). Huttonea fimbriata, (US 

266 172). 

Nigritella nigra, Kurzweil s.n. (Balogh). 

Ophrys cornuta, Reichenger s.n. (US); 0. fusca, 

Lager s.n. (US); 0. morio, Kurzweil s.n. (Balogh); 

0. rotundijolia, Dike s.n. (US); 0. ustulata, Kurzweil s.n. (Balogh); Orchis coriophora, Kurzweil s.n. 

(Balogh). 

Peristylus plantagzneus, Ramamoorthy 2008 

(US). Perularia fuscescens, Bach s.n. (US); P. ussuriensis, Matsuki 342 (US). 

Piperia unalasensis, Balogh 1056 (US). 

Platanthera sp., Greenwood 904 (AMO); P. 

nivea, Balogh 1066 (US); P. ciliaris, Zaret s.n. 

(MARY). Pterygodium catholicum, Robyns 17938 

Satyrium coriijolium, Clements s.n. (NCBG); S. 

mairei, Rock 169 13 (US); S. nepalense, Schneider 

298 1 (US). Schizochilus sandersonii, Bayliss 2541 

(US). Serapias cordigera, Teles and Rainka 331 

(US); S. paruiflora, Kurzweil s.n. (Balogh). Stenoglottis fimbriata, Lodd 6975 (US); S. longijolia, 

cultivated (Balogh). 

(US). 

Traunsteinera globosa, Kurzweil s.n. (Balogh). 

EPIDENDRo1DEAE.-Epidendroid: Acianthus 
NUMBER 61 65 

exsertus, Bernhardt s.n. (Balogh); A. fornicatus, 

(Canberra Botanical Garden); A. renqormis, 

Gunn 189 (US). Acostaea unicornuta, Luer 760 

(SEL). Adrorhiza purpurascens, Jayasuriya and 

Balasubramanium 1228 (US). Aglossorhyncha jabiensis, Dressler s.n. (Balogh). Agrostophyllum 

aseki, Dressler s.n. (Balogh); A. majus, Dressler 

s.n. (Balogh). Alamania punicea, Greenwood s.n. 

(AMO); Amblostoma tridactylum (SI 781 8). Appendicula cornuta, Dressler s.n. (Balogh). Arethusa 

bulbosa, Cheney s.n. (US). Arpophyllum sp., Hagsater 4502 (AMO), Greenwood 412 (AMO); A. 

giganteum, Suarez 17 1 (AMO). Artorima erubescens, Lamas s.n. (Balogh). Arundina graminqolia, 

Dressler s.n. (Balogh). 

Barkeria cyclotella (SI 78 1 184); B.melanocaulon, 

MacDougall s.n. (Balogh); B. vanneriana, Greenwood s.n. (AMO). Bletia sp., Greenwood 1093 

(AMO); B. ensqolia, Greenwood 1035 (AMO); B. 

liliacina, Greenwood 1094 (AMO); B. purpurea, 

Balogh 962 (US); B. reflexa, Suarez (AMO); B. 

roezlii, Balogh 744 (US); Greenwood 1033 

(AMO). Bletilla striata, cultivated (Balogh). Brassavola nodosa, cultivated (Balogh). Broughtonia 

sanguinea, (SI 78 1 196). Bulbophyllum baileyi, SH 

1138 (Balogh); B. exiguum, Clements 2308 

(NCBG); B. sp., (SI 79800). 

Cadetia sp., (SEL-77-2788). Calanthe sp., Dressler s.n. (Balogh); C. brevicornu, Dressler s.n. (Balogh); C. rubens, cultivated (Balogh). 

Calopogon multajlorus, Curtiss 2802 (US). 

Calypso borealis, Sandberg 93 (US); C. bulbosa, 

Peterson (Balogh). Cattleya sp., (SI); C. aurantica, 

Greenwood (AMO); C. bowringiana, (SI 76563); 

C. skinneri, Pollard (AMO). Ceratostylis rubra (SI 

7883). Chysis maculata, Dressler s.n. (Balogh). 

Cirrhopetalum collettii, cultivated (Balogh); C. elatum (SEL 883). Cleistes divaricata, Chapman s.n. 

(US); C. rosea, Dressler s.n. (Balogh). Coelia bella, 

Greenwood 540 (AMO); C. guatemalensis, Greenwood 658 (AMO). Coelogyne graminqolia, Dressler s.n. (Balogh); C. mayeriana, (SEL); C. pandurata (SI); C. virescens (SI). Corybas jimbriatus, 

(NCBG 761 06 1); Corybas prunosus, Bernhardt 

s.n. (Balogh). Cryptarrhena guatemalensis, Dressler 3550 (Balogh); C. lunata, Dressler s.n. (Balogh). Cryptophoranthus beloglottis, (SEL 5-75-1 l); 

C. tribuloides, Suarez (AMO). 

Dendrobium aggregatum, cultivated (Balogh); D. 

antennatum (SEL); D. bullenianum (SEL 23-74- 

561); D. delacourii (SI 76400); D. discolor, Bernhardt s.n. (Balogh); D. macbrownii, SH 1 145 (Balogh); D. mijakei (SI 792139); D. pierardii, cultivated (Balogh); D. ruppianum, Clements 2342 

(NCBG); D. secundum (SEL); D. torerensis, SH 

1 14 1 (Balogh). Dendrochilum glumaceum, cultivated (Balogh); D. insectqera, Robinson and Klass 

(US). Dichaea morrisii, Dressler s.n. (Balogh); D. 

panamense, Dressler s.n. (Balogh). Dilochia cantleyi, Dressler s.n. (Balogh). Dimeranda emarginata 

(SI 79107 1). Dresslerella pertusa, Dressler s.n. 

(SEL). 

Elleanthus sp. (SI 8 1064); E. capitatus, Dressler 

406 1 (Balogh); E. trilobatus, (SI 791 875). Encyclia 

asperula, Dressler and Pollard s.n. (AMO); E. 

citrina, Greenwood 987 (AMO); E. cochleata, Balogh s.n. (US); E. candollii, Balogh 1063 (US); E. 

livida, Balogh 795 (US); E. pentotis, Dressler s.n. 

(Balogh); E. pollardiana, Greenwood 687 

(AMO); E. pterocarpum, Hagsater s.n. (AMO); E. 

tampense, Balogh (US); E. tripunctata, Balogh s.n. 

(Balogh); E. vespa, Balogh s.n. (Balogh). Epidanthus paranthicus, Hagsater 567 1 (AMO). Epidendrum acunae, Hagsater 2073 (AMO); E. alfaroi, 

Dressler 5343 (Balogh); E. anisatum, Hagsater 

5473 (AMO); E. anoglossum, Hagsater 6344 

(AMO); E. barbeyanum, Hagsater 5643 (AMO); 

E. bqalce, Hagsater 59 10 (AMO); E. bisulcatum, 

Hagsater 6457 (AMO); E. brateosum, Hagsater 

6357 (AMO); E. brownoides (SI); E. cerinum, Hagsater 5776 (AMO); E. ciliare (SI, SEL); E. clowesii, 

Behib 702; E. cnemidophorum, Hagsater 4305 

(AMO); E. conopseum, Balogh 720 (US); E. corrqolium (Stenoglossum), Dressler s.n. (Balogh); 

E. costatum, Hagsater 4303 (AMO); E. curuicolumna, Hagsater 6520 (AMO); E. dgorme, Hagsater 4670 (AMO); E. dgusum, Hagsater 6101 

(AMO); E. eximium, Dix 200876 (Balogh); E. 

floribundum, Hagsater 5902 (AMO); E. gregorio, 

Hagsater 641 0 (AMO); E. ilense (SEL); E. incomptum, Hagsater s.n. (AMO); E. juergensenii, Hagsater 4299 (AMO); E. jejunum, Hagsater 6062 

(AMO); E. latilabrum, Hagsater 6390 (AMO); E. 

latqolium, Hagsater 5228 (AMO); E. laucheanum, 
66 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Hagsater 6 12 1 (AMO); E. longicaule, Hagsater 

6634 (AMO); E. magnzjkum, Hagsater 2600 

(AMO); E. marmoratum, Hagsater 6043 (AMO); 

E. morganii, Hagsater 6586 (AMO); E. moritzii, 

Hagsater 5075 (Balogh); E. moyabambae, Hagsater 6546 (AMO); E. myrianthum, Hagsater 5085 

(AMO); E. nagelii, Hagsater 4735 (AMO); E. 

nocturnum, Lamas s.n. (Balogh); E. nonanisatum, 

Hagsater 5887 (AMO); E. nutans, Hagsater 5208 

(AMO); E. oaxacanum, Hagsater 5753 (AMO); 

E. oerstedii; E. omissum, Greenwood s.n. (AMO); 

E. otavalo, Hagsater 59 14 (AMO); E. pachyrachis, 

Hagsater 6384 (AMO); E. patens, Behib 709 

(Balogh); E. parkinsonianum, Behib s.n. (Balogh); 

E. porpax (Neolehmannia), Dressler s.n. (Balogh); 

E. propinquuh, Hagsater s.n. (AMO); E. pugioniforme, Hagsater s.n. (AMO); E. obesum, Hagsater 

6330 (AMO); E. radicans, Balogh s.n. (US); E. 

radioferens, Hagsater 45 15 (AMO); E. ramosum, 

Hagsater 5596 (AMO); E. rowleyi, Hagsater 4827 

(US); E. sanchoi, Hagsater 6459 (AMO); E. scriptum, Hagsater 5907 (AMO); E. silacayoapan, 

Hagsater 6030 (AMO); E. skutchii, Hagsater 

6259 (AMO); E. stanifordianum, Hagsater s.n. 

(AMO); E. subviolascens, Hagsater 5778 (AMO); 

E. turialvae, Hagsater 6419 (AMO); E. vescicatum, Hagsater 6149 (AMO); E. viejii, Hagsater 

5348 (AMO). Epipogzum aphyllus, Baenitz s.n. 

(US); E. roseum, Nicholson 179 (US). Epistephium 

amplexicaule, Barriga 08293 (US), Britton and 

Rusby 2908 (US). Eria inornata, Clements 2344 

(NCBG); E. paniculata, Dressler s.n. (Missouri 

Botanical Garden). Eriaxis rigida, Buchholz 1003 

(US), McKee 3302 (US). 

Flickingeria comata, SH 1149 (Balogh). 

Galeola casythoides, Helms 11-1900 (US). Gastrodia elata, Teng 90332 (US); G. sesamoides, 

Bernhardt s.n. (Balogh). Glomera obtusa, Dressler 

s.n. (Balogh). Glossorhyncha sp., Dressler s.n. (BalHagsatera brachycolumna, McCullough 77-37 

(Balogh). Hexalectris brevicaulis, Greenwood 

1059 (AMO). Hexisea bidentata, Balogh 73 1 (Balogh). Hippeophyllum scotechini, Sinclair 565 l(US). 

Homalopetalum pumilo, Hagsater 73 (AMO). 

Zsochilus sp., Hagsater 5808 (AMO); 1. auranogh). 

tiaca, Hagsater 45 18 (AMO); I. linearis, Balogh 

7 19 (US); I. major, Montebello s.n. (Balogh). 

Jacquinella cobanensis, Balogh s.n. (Balogh); J. 

equitantijolia, Miller s.n. (Balogh); J teres, Balogh 

s.n. (Balogh). 

Koellensteinia kellneriana, Dressler s.n. (BalLaelia albida, Pollard s.n. (AMO), Balogh s.n. 

(Balogh); L. anceps, Pollard s.n. (AMO); L. autumnalis, Greenwood s.n. (AMO); L. cinnabarina (SI 

801097); L. dickinsonii, Hagsater 5708 (AMO); 

L. flavu (SI 78246); L. furfuracea, Greenwood 

s.n. (AMO); L. gouldiana, Hagsater 10-78 

(AMO); L. xanthrotropis, Moore s.n. (Balogh). 

Laeliopsis domingensis, Hagsater s.n. (AMO). Lecanorchis brachycarpa, Ohiui and Walker 7023 

(US). Lepanthes sp., Balogh 882 (Balogh). Liparis 

draculoides, Greenwood 983 (AMO); L. elata, 

Greenwood 759 (AMO); L. galeottiana, Greenwood 889 (AMO); L. lilifolia, Balogh s.n. (US); 

L. reflexa, SH 1139 (Balogh). 

Malaxis carnosa, Greenwood 717 (AMO); M. 

latifolia, SH 1143 (Balogh); M. lepidota, Greenwood 903 (AMO); M. marsupichila, Clements 

1148 (NCBG); M. soulei, Greenwood 811 

(AMO); M.fastigata, Greenwood 626 (AMO); M. 

javesiae, Greenwood 1068 (AMO); M. rosei, 

Greenwood s.n. (AMO). Masdevallia aureopurpurea (SI 79187); M. brachyura (SI) 792270); M. 

cibilis (SI 78292); M. infracta (SI); M. unzjlora, 

Balogh s.n. (Balogh). Meiracyllium trinasutum, 

Dressler s.n. (Balogh). Monophyllorchis maculeatus, Dressler s.n. (Balogh). 

Nervilia aragoana, Fosberg 37453 (US); A;. 

plicata, Saldanka and Ramamoorthy 1706 (US). 

Nidema boothiana, Balogh 802 (Balogh). 

Oberonia emarginata, King 5550 (US), 0. japonica, Makino s.n. (US). Octarrhena condensata, 

Dressler s.n. (Balogh). Octomeria graminifolia 

(SEL). Oestedella myriantha, Hagsater s.n. 

(AMO); 0. pajetense, Hagsater s.n. (AMO); 0. 

pinnifera, Hagsater 6502 (AMO); 0. ramosum, 

Hagsater s.n. (AMO); 0. schweinfurthianum, Hagsater 4351 (AMO); 0. wallisci, Tschen and Hagsater s.n. (AMO). 

Palmorchis sp., Dressler s.n. (AMO); P. trilo- 
NUMBER 61 67 

bulata, Dressler 3683 (Balogh). Phaius australis, 

Clements 2345 (NCBG). Pholidota sp. (SEL); P. 

pallida, Clements 2304 (NCBG). Phreatia robusta 

(Rhynchophreatia), Clements 2347 (NCBG). Physinga sp., Hagsater 5289 (AMO); P. physodes, 

Hagsater sen. (AMO). Physosiphon tubatus (SEL 

4 1-76-5). Physothallis harlingii (SEL 44-75-240). 

Platystele stenostachya (SEL 1 14-76-6). Pleurothallis sp., Balogh 888 (Balogh); P. caespitosa (SI 

79594); P. ghiesbrichtiana, unknown (Balogh); P. 

longissima, Balogh s.n. (Balogh). Pogonia ophioglossoides, Mearns 69 (US), Smith 205 (US). Ponera striata, Balogh s.n. (Balogh). Pseudoepidendrum spectabile, Hagsater 3390 (AMO). Psilochilus sp., Dressler s.n. (Balogh). 

Restrepia xanthophthelma (SEL 123-76-74). 

Rhyncholaelia glauca, Jackson s.n. (Balogh). 

Scaphyglottis sp. (SI); S. sp., Balogh 833 (Balogh). Scaphosepalum sardinata (SEL 30-74-207). 

Schomburgkia superbiens, Jones s.n. (Balogh); S. 

lueddemannii, Concavas s.n. (Balogh); S. tubicinis, 

Greenwood s.n. (AMO); S. wendlandii, unknown 

(Balogh). Sirhookera lanceolata, Saldanka and Ramamoorthy 562 (US); S. latijiolia, Jarrett, Saldanka, and Ramamoorthy 696 (US). Sobralia 

amabilis, Dressler s.n. (Balogh); S. decora, Dodson 

s.n. (SEL); S. fenzliana, Dressler s.n. (Balogh); S. 

fragrans (SI); S. macrantha (SI). Stenoglossum coryophorum, Dressler s.n. (Balogh). Stereosandra 

javanica, Ramos 12 124 (US), Fosberg 37285 

(US). Sunipia bicolor, Rock 7070 (US); S. racemosa, Dressler s.n. (Balogh); s. scariosa, Forrest 

26732 (US). 

Thelasis carinata, Clements 2306 (NCBG). 

Thunia alba, Troth 880 (US). Triphora trianthophora, Luer s.n. (SEL). 

Vanilla sp. (SEL); V. edwardii, Reitiz and Klein 

s.n. (US); V. pauczjlora, Dressler s.n. (Balogh); V. 

planfolia SEL. 

Wullschlaegelia aphylla, Dressler 4940 (Balogh); W. calcarata, Dressler 4646 (MO). 

Xyllosum pallidajlora, unknown (Balogh). 

Yoania japonica, Mykosan (US). 

VANDOID GROUP.-Acampe multzjlora (SEL 8 1- 

1049). Acineta blanca, Mejia s.n. (Balogh). Acriopsis javanica, cultivated (Balogh); A. nelsoniana, 

Dressler s.n. (Balogh). Ada sp., Schmidt s.n. (Balogh). Aerangis friesorum SI 81 509. Aeranthes sp., 

(SEL 78-149). Aerides mitratum (SI 7812). Angraecum distichum (SEL); A. magdulense, (SEL 78- 

380). Anguloa clftonii, Escobar s.n. (Balogh). Arachnis jlosaeris (SI). Aspasia epidendroides, Dressler s.n. (Balogh); A. principissa, Dressler s.n. 

(Balogh). 

Bollea lalindei, Mejia s.n. (Balogh). Brassia arcuigera (SEL); B. gireoudiana (SEL); B. maculata, 

(SI); B. signata, Suarez s.n. (AMO). 

Camaridium imbricatum (SI 78 1056). Campylocentrum micranthum, Foster s.n. (Balogh). Catasetum callosum, Dressler s.n. (Balogh); C. crystianum, (SEL); C. expansum, (SEL); C. integerimum, Jones s.n. (SEL); C. luridum, Dressler s.n. 

(Balogh); C. macrocarpum, Balogh s.n. (Balogh); 

C. roseum, Dressler s.n. (Balogh); C. saccatum, 

Pabst s.n. (Balogh); C. sanguineum, (SEL); C. viridijlavum, Balogh 830 (Balogh). Centropetalum 

sanguineum, (SEL). Ceratostylis rubra, (SI 7883); 

Chondrorhyncha discolor, (SI 79820). Cirrhaea saccata, (SEL). Cleisostoma macrantha, (SI 781 le50). 

Clowesia russelliana, (SEL); C. warsceswezii, Balogh s.n. (Balogh), Dressler sa. (Balogh). Cochleanthes discolor (SEL). Comparettia macroplectron 

(SI 79325). Corallorhiza striata, Balogh 977 (US). 

Coryanthes maculata, Dressler s.n. (Balogh). Cryptarrhena lunata, Dressler s.n. (Balogh). Cycnoches 

chlorochilon, Balogh 877 (Balogh); C. diandre, 

Dressler s.n. (Balogh); C. egertonianum (SEL); C. 

loddigessii (SEL); C. ventrycossum, Dressler s.n. 

(Balogh); C. warscewiczii, Suarez s.n. (Balogh); C. 

sp., Suarez 391 (AMO). Cymbidium sp. (SI); C. 

caniculatum, SH 1144 (NCBG); C. eburneum, (SI 

79840). Cyrtochilum carderi, Posada s.n. (Balogh). 

Cyrtopodium punctatum (SI). Cyrtorchis monteirae 

(SI 792 172). 

Dichaea morrisii, Dressler s.n. (Balogh); D. panamensis, Dressler s.n. (Balogh). Dipodium punctatum, Clements 2272 (NCBG); D. pandanum, 

Dressler s.n. (Balogh). Doritis pulcherrima (SEL). 

Dressleria helleri, Balogh 5.n. (Balogh). 

Eriopsis wercklei, Dressler 3940 (Balogh). Erycina echinata, Dressler s.n. (Balogh). Eulophia faireana, Dressler s.n. (Balogh); E. petersii, unknown 
68 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

(Balogh); E. tuberosa, Dressler s.n. (Heidelberg). 

Fernandezia hartwegii, Dressler s.n. (Balogh); 

F. sanguinea, Dressler s.n. (Balogh). 

Galeandra devonianum (SI). Gastrochilus bellinus (Frankfurt). Genyorchis pumila (Utrecht). Geodorum sp., Dressler s.n. (Balogh). Gomesa crispa 

(SI 78220). Gomeza planifolia (SI 792196). Gongora claviodora, Dressler s.n. (Balogh); G. gratulabunda, Mejia s.n. (Balogh); G. sp., Suarez 169 

(AMO); G. tricolor (SEL 1-1975-618); G. truncata 

(SEL 105-76-25). Govenia bella, Greenwood 956 

(AMO); G. capitata, Greenwood 873 (AMO); G. 

lagenophora, Greenwood 4 17 (AMO); G. liliacea, 

Greenwood 556 (AMO); G. mutica, Greenwood 

574 (AMO); G. superba, Greenwood 902 (AMO); 

G. tequiliana, Dressler and Hagsater 27 15 

(AMO). 

Haraella odorata (SI). Houlletia tigrina, Mejia 

s.n. (Balogh). Huntleya burtii, Balogh s.n. (BalZonopsis utricularioides, Dressler s.n. (Balogh). 

Jumellea sagittata (SI 78234). 

Kefersteinia paruilabris, Mejia s.n. (Balogh). Kegeliella kepperi, Dressler s.n. (Balogh). Koellensteinia kellneriana, Dressler s.n. (Balogh). 

Listrostachys pertusa, Dressler s.n. (Heidelberg). Lockhartia micrantha, Balogh 58 1 (Balogh); 

L. oestedii, Dressler s.n. (Balogh); L. pallida, Dressler s.n. (Balogh). Lueddmannia pescatorei, Mejia 

s.n. (Balogh). Lycaste aromatica (SI 8 1 1874); L. 

consobrina (SI 76960); L. cruenta, Hagsater s.n. 

(AMO); L. deppei, Suarez s.n. (AMO); L. leucantha (SI); L. powellii, Dressler 3867 (Balogh). 

Masdevallia asperrima (SI 8 12667). Maxillaria 

brunnea, Dressler s.n. (Balogh); M. juergensii (SI 

77 1 18); M. rufescens, Balogh s.n. (Balogh). Mexicoa ghiestbrechtiana, Balogh 970 (Balogh). Miltonia “Anne Warne” (SEL); M. bluntii (SI); M. 

jlavescens (SI 78319); M. spectabilis (SI 78321-A). 

Miltoniopsis roezlii, Dressler 4084 (MO). Mormodes cartonii, Balogh 8 1 1 (Balogh). Mormolyca 

ringens, Dressler s.n. (Balogh). 

Neobenthamia (S1 76404). Notylia maculilabris, 

Dressler 1096 (MO); N. trisepala, (SI 7833 1). 

Oberonia emarginata, King 5550 (US). Odontoglossum sp., Hagsater 43109 (AMO); 0. apterum, 

ogh). 

Greenwood s.n. (AMO); 0. cordatum, Halbinger 

s.n. (Balogh); 0. harryanum, Dressler s.n. (Balogh); 0. maculatum, Balogh s.n. (Balogh); 0. majace, Hagsater 5341 (AMO); 0. pulchellum Balogh s.n. (Balogh); 0. reichenheimii, Balogh 970 

(US). Oeceoclades maculata, Foster s.n. (Balogh). 

Oncidium aurisassinarum (SI 797005); 0. desertorum, Stern 216 (Balogh); 0. durangensis, Hagsater 3769 (AMO); 0. fasciculatum, Hagsater 5332 

(AMO); 0. leucochilum, (SEL); 0. maculatum, Pollard s.n. (AMO); 0. oblongatum, Hagsater 3364 

(AMO); 0. obryzatum, Hagsater 4177 (AMO); 0. 

ornithorrhynchum, Pollard s.n. (AMO); 0. papilio 

(SI); 0. rejlexum, Balogh 789 (Balogh); 0. stenotis 

(SI 7625 1). Ornithocephalus bicornis, Balogh 840 

(Balogh); 0. cochlearueformis, Balogh s.n. (Balogh); 0. tripterus, Dressler s.n. (Balogh). Ornithophora radicans, Dressler s.n. (Balogh). Osmoglossum egertonii, Hagsater 6518 (AMO); 0. pulchellum, Pollard s.n. (AMO). 

Pachyphyllum distichum, Hutchison 6655 (US); 

P. pasti, Holmgren s.n. (US); Paradisianthus micranthus (SEL); Peristeria elata, Balogh s.n. (Balogh). Phalaenopsis corni-cervi (SI 78 1076); P. luemanniana, Brecht 1969 (Balogh). Plectrophora 

data, JFM 882 (Balogh); P. cultifolia (SI 78 1440). 

Polycynis gratiola, unknown (Balogh); P. adansonae (SI 792259); P. bella (Missouri Botanical 

Garden). Polystachya concreta (SI 7774); P. galangoreae (SI); P. tessellata (SI 792260). 

Rangaeris amaniensis, Dressler s.n. (Balogh). 

Renanthera imschootiana, Dressler s.n. (Balogh). 

Rhynchostylis retusa, Dressler s.n. (Balogh). Rodriguezia granadensis, Dressler s.n. (Balogh). Rossioglossum schlieperianum, Hagsater 6433 (AMO); 

R. splendens, Hagsater 3893 (AMO); R. williamsianum, Hagsater 6630 (AMO). 

Sarcanthus pachyphyllus (SI 792 146). Schomburgkia tubicinis (SEL). Sievekingia burcheri, Dressler s.n. (Balogh); s. columbiana, Dressler s.n. 

(Balogh). Sigmatosalix guatemalensis, Lent s.n. 

(Balogh); S. mexicana, Hagsater 4772 (AMO); S. 

sp. Balogh 814 (Balogh). Stanhopea connata (SI 

79 1282); S. costaricensis, Dressler s.n. (Balogh); 

S. ecornuta, Dressler 4092 (Balogh); S. guttulata 

(SI); S. oculata, Suarez 82 (AMO); S. tricornis, 
NUMBER 61 69 

Dressler s.n. (Balogh); S. tigrina, Dressler s.n. 

(Balogh); S. wardii, Dressler s.n. (Balogh). Stelis 

sp., Balogh 893 (Balogh). Stenocoryne racemosa 

(SI 78477); S. vitellina, Dressler s.n. (Balogh). 

Telipogon klotscheanus, Dressler s.n. (Balogh). 

Teuscheria dodsonii, Dressler s.n. (Balogh). Thecostele alata, Dressler s.n. (Balogh); T. maingayi, 

Dressler s.n. (Balogh). Thrixospermum elongatum 

(SI 79 1401). Tipularia discolor, Balogh 908 (US). 

Trichocentrum bursigera, unknown (Balogh); T. 

pfavii, Balogh s.n. (Balogh); T. tigrinum (SEL). 

Trichoglottis fasciata (SEL). Trichopilia elegans (SI 

78486); T. galeottiana (SI 78487); T. hennisiana, 

Mejia s.n. (Balogh). Trigonidium egertonianum, 

Pollard s.n. (AMO). Trizeuxis falcata (SI 792008). 

Vanda coerulescens (SI 78 146 1); V. sp. (SI); V. 

limbata (SEL 26-75-3). Vandopsis lissochiloides 

(SEL). 

Warcella cyanea, Mejia s.n. (Balogh). Warmingia sp. (SEL); W. elegans, Dressler s.n. (Balogh). 

Warscewiczella velata, Lasca s.n. (Balogh). 

Xylobium colleyi, Dressler 3748 (Balogh); X. corrugatum, Mejia s.n. (Balogh); X. elongatum (SI 

79 1004); X. foveatum (SI 79 1005). 

Zygopetalum mackey (SI 76956). 
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72 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 1 .-Floral morphology of Orchidaceae, whole flowers: a, Neuwiedia veratifolia Blume, 

photo by J.B. Comber; b, Apostasia wallichii R. Brown, J.B. Comber: c, Cypripedium irapeanum 

Llave and Lexarza, E.W. Greenwood: d, Paphiopedilum venustum Pfitzer, P. Burns-Balogh: e, 

Paphiopedilum spicereanum Pfitzer, P. Burns-Balogh:f, Diuris longifolia R. Brown, M.A. Clements; g, Prasophyllum parvifolium Lindley, M.A. Clements; h, Cryptostylis javanica J.J. Smith, J.B. 

Comber; i, Anoectochilus setaceus Lindley, J.B. Comber; j, Erythrodes sp., E.W. Greenwood: k, 

Ponthieva tuerkheimii Schlechter, E.W. Greenwood; 1, Caladenia patersonii R. Brown, M.A. 

Clements; m, Thelymitra nuda R. Brown, M.A. Clements; n, Platanthera ciliaris Lindley, P. 

Burns-Balogh: 0, Habenaria koordersii J.J. Smith, J.B. Comber: p, Habenaria distans Grisebach, 

E.W. Greenwood. 
NUMBER 61 73 
74 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 2.-Floral morphology of Orchidaceae, whole flowers: a, Galearis spectabilis Rafinesque, 

P. Burns-Balogh; b, Disperis javanica J.J. Smith, J.B. Comber; c, Arundina graminfolia Hochinger, J.B. Comber: d, Bletia sp., E.\V. Greenwood; e, Hexalectris brevicaulis L.O. Williams, E.W. 

Greenwood; J; Pleurothallis sp., E.W. Greenwood; g, Stelis sp., P. Burns-Balogh: h, Mormodes 

ignea Lindley and Paxton, P. Burns-Balogh; i, Oncidium sp., E.W. Greenwood; j, Clowesia 

thylaciochila (Lenioire) Dodson, E.W. Greenwood; k, Clowesia warscewitrii (Lindley and Paxton) 

Dodson, P. Burns-Balogh: 1, Cynoches warscewitrii Reichenbach filius, E.W. Greenwood: m, 

Catasetum sanguineurn Lindley (O), P. Burns-Balogh; n, Catasetum sanguineum Lindley (a), P. 

Burns-Balogh. 
NUMBER 61 75 
76 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE J.-Floral morphology of Orchidaceae, dissected parts of flowers: a, Cypripedium acaule 

Aiton, P. Burns-Balogh; b, Sarcoglottis gutturosa (Reichenbach filius) Ames, E.W. Greenwood; 

c-J Sarcoglottis paucijlora (Richard and Galeotti) Schlechter, E.W. Greenwood; g,h, Ponthieva 

tuerckheimii Schlechter, E.W. Greenwood; i, Ponthieva racemosa (Walter) Mohr, E.W. Greenwood; j, Rhizanthella gardneri Rogers, P. Burns-Balogh; k, Habenaria distans Grisebach, E. W. 

Greenwood; 1, Bletia ensvolia L.O. Williams, E.W. Greenwood; m, Hexalectris brevicaulis L.O. 

Williams, E.W. Greenwood; n, Pleurothallis tribuloides (Swartz) Lindley, E.W. Greenwood. 
NUMBER 61 77 
78 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 4.-Floral morphology of Orchidaceae, dissected parts of flowers: a, Lepanthes sp., P. 

Burns-Balogh; 6, Oncidium sp., E.W. Greenwood; c, Clowesia thylaciochila (Lernoire) Dodson, 

E.M’. Greenwood; d, Cycnoches warscewitzii Reichenbach filius, E.W. Greenwood; e-g, Gongora 

truncata Lindley, E.M’. Greenwood; h-k, Stanhopea oculata (Loddigues) Lindley, E.W. Greenwood. 
NUMBER 61 79