DNA DATA AND ORCHIDACEAE SYSTEMATICS :

A NEW PHYLOGENETIC CLASSIFICATION.

Mark W. Chase

Jodrell Laboratory ,Royal Botanic Gardens ,Kew, Richmond , Surrey TW9 3DS, UK .

Kenneth M.Cameron

The Lewis B. and Dorothy Cullman Program for Molecular Systematics Studies,The New York Botanical Garden, Bronx, New York 10458-5126 , USA.

Russell L. Barrett

Kings Park and Botanic Gardens and Parks Authority ,West Perth , 6005,Western Australia and Plant Biology, Faculty of Natural and Agricultural Sciences ,The University of Western Australia ,Crawley 6009, Western Australia .

John V. Freudenstein

Ohio State University Herbarium , 1315 Kinnear Road ,Columbus , Ohio43212-1157,USA.

Orchidaceae are rapidly becoming one of the best-studied families of the angiosperms in terms of infra-familial phylogenetic relationships. These studies demonstrate that several previous concepts about phylogenertic patterns werer incorrect , which make all previous classifications in need of review . Therefore , in this paper we describe the emerging patterns and propose a new phylogenetic classification of Orchidaceae that accords with these newly discovered relationships. We recognise five subfamilies: Apstasioideae ,Vanilloideae ,Cypripedioideae, Orhidoideae are sister to all the rest , followed successively by Vanilloideae ,Cypripedioideae and the remainder of the monandrous orchids,Orchidoideae and Epidendroideae . Although only an interim classification , it should help to focus other areas of orchid research and stimulate the creation of new hypotheses that will direct orchid researchers to new questions .

1. Introduction

For many years , orchid classification has been based almost exclusively on features of their gymnostemium or column (Brown , 1810; Lindley , 1840 ; pfitzer , 1887; schlechter , 1926 ; Swarts, 1800). In the two most recent of these systems , an evolutionary progression was hypothesised from two or three anthers in the apostasioid orchids (apostasia and Neuwiedia )through two in the cypripedioids (Cyprepedium, Medipedium, Paphiopedilum , Phragmipedium , and Selenipedium) to once in the monandrous orchids (Epidendroideae , Orchidoideae and Spirnthoideae sensu Dressler, 1993). Within the monandrous orchids ,which contain the great majority of orchid taxa ,classification has depended largely on whether pollen in the anther was loose or formed into packets of various sorts, including hard pollinia . In the apostasioids , pollen is powdery as it is in most groups of Asparagales (sensu Angiosperm Phylogeny Group (APG), 1998), but in all other orchids , pollen is at least sticky and self-adherent so that it travels in packets , which is probably related to the large number of ovules in the ovaries of most orchids .In the most highly evolved groups of epidendroid orchids (roughly 80% of the species in the family ; Dressler , 1993), pollen is firmly bound into hard pollinia deposited as complete units in the stigmatic cavity , but in the other monandrous orchids, there is every possible intermediate stage between free monads and hard pollinia . Most systems have also emphasised the other structures that comprise pollinaria ,such as stipes ,caudicles ,and viscidia ,but only a few older classifications (e.g.Pfitzer, 1887)have incorporated any number of vegetative characters.

Because orchid classification has largely been based on the relative degree of organisation of the pollinia , the distinction between Neottioideae and Epindendroideae has been highly problematic , such that the more primitive group , Neottioideae , has been variously narrowly and broadly defined . In Dressler's two schemes (1981; 1993) ,the neottioid orchids were narrowly treated . In addition to circumscription of the neottioids , the other major group of orchids that has been problamatic is the vanilloids .Their columns are much like those of the epidendroids , but vegetatively they are highly divergent from all other orchids (Cameron and Dickison, 1998; Stern and Judd , 2000).

More recently , orchid systematists have begun the process of incorporating other categories of morphological information into their classifications (Dressler and Dodson, 1960 ; Garay, 1960; 1972;Vermeulen , 196; Rasmussen , 191985; Burns -Balogh and Funk , 1986 ; Brieger , Butzin and Senghas, 1995; Szlachetko , 1995), but his process has only infrequently been couched in terms Senghas , 1995; Szlachetko, 1995), but this process has only infrequently been cauched in terms of explicitly phylogenetic studes (Freudenstein and Rasmussen , 1999). Burns -Balog and Fuck (1986) presented their arguements om cladogram format, but no formal analysis was conducted . Dressler(1981; 1993) also conveyed his ideas about relationships in the form of cladograms with characters mapped onto them , but their structure was purely intuitive . The results of the morphological analyses og Frudention and Rasmussen (1999) indicated that the high degree of hierarchial structure in all previous classifications of Orchidaceae was not warranted ; this assertion at lower taxonomic levels. They did , in contrast , provide support for some of the various subfamilial groupings recognised in most previous systems of classification , such as Apostasioideae, Cypripediodeae , Orchidoideae and Epidendroideae.

Molecular data have come to play an increasingly important role in angiosperm calssification (Chase et al., 1993; 2000 a ; b ; APG, 1998; Soltis , Soltis and Chase, 1999;Chase, Fay and Savolainen , 2000 ; Savolainen et al ., 2000; Soltis et al ., 2000), and although the main focus has been at the supra-familial level, increasingly efforts are being focused on familial classification (Sheahan and Chase, 1996; 2000; Chase et al ., 2000 c; Richardson , Fay and Chase , 2000 ). Within Orchidaceae, numorous DNA phylogenetic studies have now been published , ranging from the whole family (Neyland and Urbatsch, 1993; Chase et al ., 1994 ; Cameron et al ., 1999; Molvray , Kores and Chase , 2000 ; Freudenstein , Senyo and Chase, 2000 a ; b) , subfamilies (Cox et al ., 1997; Kores et al ., 1997), tribes (Cameron and Chase , 1999; Douzery et al ., 1999; Kores et al., 2000 ; Whitten ,Williams and Chase , 2000; Goldman et al ., 2001), subtribes (Chase and Palmer , 1989; 1992; 1997; Chase and Hills , 1992; Yukawa , Cameron and Chase , 1996; Pridgeon et al ., 1997; Pridgeon and Chase , 1998 ; van den Berg et al ., 2000 ; williams et al ., 2001) . In terms of general patterns of orchid relationships, these studies have portrayed a remarkably similar result , regardless of which genome they represent . Orchids have now been the focus of more published DNA phylogenetic studies than any other family in the angiosperms , and these have formed the basis of the on-going Genera Orchidacearum project (Prideon et al ., 1999; 2001 ; 2003), which will place future orchid research on a solid footing once it has been completed.

The position of the orchids among the lilioid monocots has also been the subject of a great deal of molecular systematic study (Chase et al ., 1993 ; 1994 ; 1995 ; 2000 c ; Chase ,Rudall and Conran , 1996), and these studies have demonstrated that the orchids are a member of Asparagales , nearest to families like Asteliaceae , Blandfordiaceae , Boryaceae ,Hypoxidaceae and Lanariaceae , with which they share a general syndrome of characters , such as inferior ovaries , simultaneous microsporogenesis and septal nectaries (although orchids exhibit both sepal and non-septal or perigonal nectaries ). Orchids are also well known for their mycorrhizal relationships ,which are also documented in Boryaceae (two Australian genera, Alania and Borya) , although the exact mycorrhizal mechanism and the type of fungi associated with these speceis are unstudied thus far (J.Conran , pers.comm.).

The overall biogeography and phylogenetic patterns of Orchidaceae are highly consistant with the family being an old one broadly distributed before the break-up of Pangaea ,such that the orchids are roughly 100 million years old (Chase , 2001). Long-distance dispersal is undoubtedly a factor in the current distribution of orchids, but their basic underlying distribution is that of a widespread , largely tropical family in which biogeography has been greatly affected by plate tectonics . Determining where such a family originated is highly speculative . They clearly could not have originated in Indomalaysia as hypothesised byGaray (1972) because that area did not exist as such at the time orchids are likely to have evolved . The common ancestor of orchids and other Asparagales was undoubtedly terrestrial , and the hypothesis that they could have been otherwise (Robin adn Burogh , 1982) was based on a spurious set of ideas about the adaptive nature of various orchid apomorphies , such as resupinatte flowers and micorspermy (for opposing atguments , see Ackermin , 1983; Bensing and Atwoood , 1984). Orchids clearly have a set traits that has pre-adapted them for an epiphytic existence , and several predominantly terrestrial lineages have species that have become epiphytic Independtly (Phaphiopedilm exul of Cypripedioideae and Lankesterella of Spiranthinae, to mention just two examples. )

2. Classification

At present , the most widely used orchid classification is that of Dressler (1993), but it and the more recent one of Szlachtko(1995) are clearly out-of -date relative to the cladistic analyses of Neyland and Urbatsch (1993) , Chase et al . (1994) , Cameron et al. (1999) and Freudenstein and Rasmussen (1999) . Therefore a new classification of Orchidaceae is desirable, and at least a new subfamilial classification is justified on the basis of the previously published evidence , both molecular and morphological . Two subfamilies are easily agreed upon by all researchers ,Apostasioideae and Cypripedioideae. A third ,Vanilloideae , has been argued upon by Szlachtko (1995), Cameron et al. (1999) , and Cameron and Chase (1999). This entity is relatively unknown to most botanists , but it is supported by a number of lines of evidence , most clearly the DNA data.

The position and distinctiveness of Spiranthoideae sensu Dressler (1993) has been problematic. Vegeratively they are like Orchidoideae in their spirally arranged leaves without sclerenchyma bundles (except in Tropidieae ) and seed testa with intercellular spaces, but their Freudenstein and Rasmussen (1999) found them to be embedded within orchidoid taxa in their morphological analysis , as have all DNA sequence data studies (Chase et al ., 1994 ; Kores et al., 1997; 2000 ; Cameron et al ., 1999). Tropidieae , which have similar terminal anthers , always fall with molecular data in the epidendroid group of genera as they did in the morphological analysis of Freudenstein and Rasmussen (1999). Seed characters show a lack of distributional correspondence with these anther and root characters ,such that the spiranthoid seed-type is found in Diurideae and Pterostylidinae but not in Tropidieae , which turn out to be epidendroid (see below) . Thus most of the evidence demonstrates that Spiranthoideae must be abandoned and submerged in Orchidoideae and Tropidieae should be transferred to Epidendroideae.

Finally , we come to Neottioideae , which as a concept have never consisted of more than a group of orchids lacking well-developed pollinia .Dressler (1981; 1993) considered these orchids to be no more than a tribe and greatly reduced the number of included genera. All phylogenetic analyses , molecular and morphological , have demonstrated that their continued recognition as a formal subfamilial grouping is unjustified . In their vegetative traits thery are clearly members of either the orchidoid or epidenroid groups , although their flowers are heterogeneous , and some have columns , such as Tropidieae , that are " spiranthoid ". If familial status for the neottioids can be discounted , then this leaves us with five subfamilies on which there is broad agreement : Apostasioideae , Cypripoideae , Epidendroideae , Orchidoideae and Vanilliodeae . The interrelationships of these five clades have not been satisfactorily resolved until recently , but a combined analysis of atp B , rbc L, mat K , psa B , trnL-F (all five plastid , the last largely non-coding ), 26S r DNA (nuclear) , and nad 1 intron (mitochondrial non-coding) strongly supports the following pattern: (Apostasioideae [vanilloideae[Cypripedioideae [Epidendroideae, Orchidoideae]]]](Fig . 1 ; Cameron , 2001 ; Freudenstein et al., 2001). This set of relashionships has important implications for the evolutionary interpretation of how the orchid column has evolved (considered below).

Within Orchidoideae s.l., most relationships are now fairly clear , as is also true in Vanilloideae , both on the basis of DNA sequences analyses . Epidendroideae are still problematic , simply due to the low levels of sequence divergence relative to the number of taxa involved , but even within these some clear patterns are emerging . Within Epidendroideae , the DNA sequence tree strongly contradicts all previous classifications , but some of the results of morphological analyses (Freudenstein and Rasmussen, 1999 ) anticipated these DNA patterns. As pointed out by Freudenstein and Rasmussen (1999) , the relationships reflected in morphology are weak hypotheses , but they too, as mentioned above , contradict most, but not all , taxonomic categories in previous classifications (for example , morphology and DNA sequences both support Dressler's , 1981,concept of vandiodeae; see below ). What we are left with is a clear impression that a new paucity compared to the numbers of taxa to be classified . This leaves us with the choice of either making no changes to the status quo or basing a new classification on patterns revealed largely by DNA sequence studies . The first choice appers unacceptable to us ; it makes no sense to continue to use any classification that has been refuted and we know to be unreliable . Scientists in other fields use classification to provide a framework for their research , and we know that large parts of the most recent orchid classification s (Dressler , 1993; Szlachtko, 1995) are erroneous and so would be a misleading basis of other kinds of studies.

Basing a classification soley on DNA data would appear to be imprudent and undesirable to many botanists. First of all, the newly circumscribed taxa will be likely to lack clear,defining characters in many cases, but that same criticism can be made aganist systems , particularly for groups circumscribed by the lack of apomorphies . For example , many epidendroid tribes , such as Arethuseae sensu Dressler (1993) are glossly polyphyletic (van de Berg et al., 2000 ; Goldman et al ., 2001). In our opinion , it is better to produce a revised classification based at first at first on DNA data alone because such a system can be " refuted" by the collection of additional data and thus be improved , whereas we are certain on the basis of current information that the previous systems are not only incorrect in many deatails but also irrefutable because they were so heavily dependent on their author's intuition . Even those that were expressed in cladogram format (Dressler , 1981; 1993; Burns-Balogh and Funk , 1986) cannot be duplicated with morphological (Freudenstein and Rasmussen , 1999)or molecular analyses (Cameron et al., 1999).

Production of a highly detailed and hierarchical classification at this time would be premature , so we have shown several subtribes without tribal affiliation . In Fig . 1 , we have also left many tribes without specific relationships , but there are several subtribes and tribes for which clear pattern have been established in studies using multiple DNA regions . We will comment below in more detail on these patterns.

2.1 Vanilloideae

The existence of five major lineages that may be recognised as subfamilies , Apostasioideae , Cypripedioideae , Epidendroideae , Orchidoideae and Vanilloideae , was noted by Chase et al. (1994), the first molecular study to include all major lineages ; Neyland and Urbatsch (1993) was published earlier , but did not sample all groups . Vanilla and its relatives have had a long history of recognition as some form of higher category (even as a family , Lindley , 1836; Garay, 1986), but possession of a single anther led to their recognition as at best a subgroup of monadrous orchids. Their single , incumbent anther and poorly organised pollinia have generally resulted in the vanilloids being lumbed with either primitive Epidendroideae or broader circumsriptions of Neottioideae , but the DNA results indicate that instead the former is based on parallelism and the latter a symplesiomorphy . Their crustose seeds have been compared with those of Curculigo (Hypoxidaceae ) and Apostasia (Garay , 1960) , but because they were viewed as relatively advanced (monandrous ) orchids ,this condition was viewed as a reversal from dust seeds. However , a position near the base of the family could mean that this seed type is plesiomorphic for orchids because it is shared with their outgroups. The seed characteristic of Apostasia and Neuwiedia are also highly aberrant relative to the dust-like seeds of most orchids,which indicates that their condition is autapomorphic and irrelevant to interpretations of orchid seed evolution .The seeds of the vanilloids therefore assume a much greater significance than previously hypothesised . Other unusual vanilloid features, such as reticulate leaf venation , are clearly highly derived and atypical in the context of Asparagales as well as orchids. Also includes in Vanilloideae are Pogoniinae, which Dressler (1993) listed as incertae sedis , although he earlier (1981) placed them with Vanillieae .

2.2 Orchidoideae

The polyphyly of Orchidoideae (if they included Diurideae and Chloraeinae ) and the embedding of Spiranthoideae in Orchidoideae were also features of the earliest molecular studies (Chase et al., 1994; Kores et al., 1997; Cameron et al ., 1999), as was the removal of Tropidieae and Diceratosteleae from Spiranthoideae to Epidendroideae . These patterns were also clear in the morphological study of Freudenstein and Rasmussen (1999).

Diseae sensu Dressler (1993) appear to be paraphyletic to Orchieae (Douzery et al ., 1998; Kores et al., 2000), which necessitates either recognition of several new small tribes or enlargement of Orchideae to include all Diseae . We favour the recognition of fewer , larger tribes , and our expansion of Orchideae to include Diseae is consistent with that philosophy. Codonorchis (formerly Chloraeinae sensu Dressler , 1993) falls in a highly divergent position as sister to Orchideae s.l . and could be included in the latter , but for the present recognition of Codonorchideae appears appropriate (Cribb and Kores, 1999).

The recognition of numerous, small subtribes within Diurideae has a historical precedent (i.e. Dressler , 1993), but such narrow delimitation has not been paralleled in Cranichideae and Orchideae. We recognise fewer subtribes here than more recent treatment (Clements and Jones , 2001;Clements et al ., 2001 ; Kores et al ., 2001) and believe that further amalgamation is desirable . The transfer of Pterostylidinae and Chloraeinae from Diurideae to Cranichideae is also well supported (Kores et al., 2000). We include Pachyplectron in Goodyerinae rather than its own subtribe because it is sister to all of the core spiranthoides (Spiranthinae and Crachidinae ;Salazar et al ., 2003).

Cranichideae sensu Dressler (1993) appear to the monophyletic but should also include Chloraeinae and Pterostylidinae ; thus Cranichideae as here circumscribed are composed of six subtribes ; Chloraeinae ,Cranichidinae , Goodyerinae , Manniellinae ,Spiranthinae and Pterostylidinae. Generic limits in Spiranthinae have been problematic , with some authors recognising a single genus drastically on their circumscription (Balogh , 1982;Garay , 1982; Szlachetko, 1995). Recent DNA-based studies on the subtribe have shown that it would be monophyletic if Galeottiella (monotypic from Mexico ) is removed to its own new subtribe (Salazar , 2003). Extensive generic recircumscription is also necessary to make the units identified in the DNA sequence study monophyletic (Salazar et al., 2003); these redefined genera are included in the Appendix .

Cranichidinae as circumscribed by Dressler (1981) are monophyletic , but Prescottinae as defined in Dressler (1993)are paraphyletic to them , so we recognise here the earlier , broader limits in which these two subtribes are combined . Goodyerinae (including Pachyplectron) require much addition study of generic limits , but the subtribe is monophyletic (Kores et al., 2001; Salazar et al., 2003).

2.3 Epidendroideae

The bulk of the monodrous orchids are included here . Several of the more phylogenetically derived and morphologically advanced tribes and subtribes are monophyletic and well studied (see below), as are the tribes forming successive sister groups to the rest at the three msot basal nodes , but the bulk of the groups falling between these two sets of tribes are not yet well characterised . The major problem here is that the number of variable positions in most of the commonly sequenced DNA regions is too low to provide resasonable estimates of phylogenetic patterns . If sequence divergence is at all clock-like within these taxa , then the reason for this lack of clear patterns would be a rapid radiation of lineages within a short period of time (for example ,see Cameron et al., 1999, for the short branch lengths along the spine of the rbcL tree in Epidendroideae ).We provide here a number of newly circumscribed tribes differing extensively from tribes with the same names in Dressler (1993) , but we admit that placement of many genera is tentative (those genera for which no DNA information is available are marked with in the Appendix ).Genera for which there is DNA information but placement is not well supported (by the bootstrap or jackknife ) are enclosed in square brackets , indicating that this requires further study . Although revising the classification of Dressler (1993) for these tribes and subtribes appears premature , we believe that a classification based on the emerging patterns from the DNA studies is an improvement over one that has largely been refuted .For example , our dismemberment and reconstitution (see below ) of Arethuseae sensu Dressler (1993)appears to be an appropriate , if only intermediate ,advancement that can serve a useful purpose of facilitating study by other orchid researchers.

The first several successive sister groups to the rest of Epidendroideae (fig.1) are all small . Neottieae as circumscribed here are the same as in Dressler (1993) , but with the addition of Palmorchis .Except for Aphyllorchis and Palmorchis , this small group of six genera are from the temperate zone of the Northern Hemisphere. Neottia is embedded within Listera , but the former is the older name. Sobralieae are likewise a small tribe of just four genera , but these are confined to the tropics of the New World, whereas Tropidieae probably have only two Pantropical genera. Most of the members of these tribes are plesiomorphic in habit (Dressler , 1981; 1993; Chase , 2001), but in terms of floral characteristics , they are more advanced , particularly Sobralieae , which have often been placed with Epidendreae (Dressler ,1993). Sobralieae have floral characteristics that parallel those of Cranichideae , in which they have often been placed (Dressler , 1981; 1993). Sobralieae and Tropidieae are sister clades in the trees placed (Dressler, 1981; 1993). Sobralieae and Tropidieae are sister clades in the trees produced by the combined DNA sequence matrix , and if this should turn out to be their final placement , then perhaps combining them into one tribe would be appropriate since they are both so small and similar vegetatively.

Circumscription of the next nine tribes/subtribes is highly problematic .Most of these are largely tropical , Old World groups for which little material has been available for inclusion in the DNA sequence studies. Dendrobiinae and Bulbophyllinae are circumsribed as Dressler (1993), except that Pseuderia has been moved to Podochiliae (Yukawa et al., 1996). Likewise , we are advocating no changes for Malaxideae relative to Dressler (1993). Nervilieae include Nervilia, plus tentatively the genera of Epipogiinae (Silvorchis, Stereosandra, and Epipogium; Molvary et al., 2000) and xerorchis (Cameron et al., 1999; Cameron , 2001).Triphoreae include the three genera placed there by Dressler (1993) plus Diceratostele, which has previosly been included in its own subtribe of uncertain placement (Dressler , 1993), but we place it here in Triphoreae on the basis of its position in the rbcL tree of Cameron et al .(1999).

Our circumscription of the heterotrophic tribe Gastrodieae is much narrower than that of Dressler (1993) and includes only Gastrodiinae ; Epipogiinae are removed to Nervilieae (see above ), and Wullschlaegeliinae (monogeneric ; Wullschlaegelia ) are included in Calypsoeae (see below). These plants are problematic for molecular systematic studies because they apparently lack many of the plastid DNA regions relied upon in most previous research (Cameron et al ., 1999; Cameron , 2001 ; Goldman et al ., 2001; Freudenstein et al., 2001). Molvray et al . (2000), however , used sequences of nuclear 18S ribosomal DNA to evaluate the placement of these genera, finding that only Gastrodiinae were monophyletic .Her study was hapered by the generally low levels of sequence of divergence for 18S rDNA sequences within orchids, which limited the number of photosynthetic orchids that could be sampled . We hope that more extensive taxonomic sampling of both the nuclear ITS non-coding rDNA as well as a portion of nuclear 26S rDNA (Freudenstein et al., in prep.), both of which have many more variable sites , will result in a more satisfactory estimate of their relationships.

Two of the most problematic tribes in terms of their taxonomic histories are Arethuseae and Calypsoeae. Both of these are treated were very narrowly circumscribed in Dressler (1981; only Calypso and Yoania ), whereas his latter treatment was much broader and more similar to that adopted here (i.e including Corallorhizinae , plus Govenia ; van den Berg et al ., 2000; Freudenstein et al ., 2001; Goldman et al ., 2001). Freudenstein (1994) published a morphological cladistic analysis of Calypsoeae (considered at that time to be Corallorhizinae), which showed a similar pattern of relationships to those found in Goldman et al . (2001).

In contrast to Calypsoeae ,Arethuseae are much more narrowly circumscribed than in Dressler (1981, 1993), and many of the excluded genera have been moved to either Collabiinae or Epidendreae (see below). Goldman et al. (2001)focused on Arethuseae and found that some of these genera (including the type genus , Arethusa ) are closetto Coelogyninae , a result also found by Cameron et al. (1999) ; van den Berg et al .(2000) and Cameron , (2001) , so here we tentatively place Arethusinae and Coelogyninae together in Arethuseae. We have some confidence that Arethusinae contain at least Anthogonium ,Arethusa , Eleorchis , Calopogon and probably Arundina . In turn, Coelogyninae are composed of Bletilla , Coelogyne (and its close relatives , such as Dendrochilum and Entomophobia ;Gravendeel et al ., 2001), Dilochia , Glomera , Thunia and Pleione . In contrast , many of the genera that Dressler (1993) placed in Arethuseae fall elsewhere . Bletia and Hexalectris (also Arethuseae sensu Dressler (1993) are strongly supported as members of Epidendreae.

Acanthephippium, Calanthe, Phaius and Spathoglottis (all Arethuseae sensu Dressler, 1993) come together with collabium , and these are here recognised as Collabiinae . Collabiinae also include also include the genera placed there by Dressler (1993; Chrysoglossum, Collabium, and Diglyphosa). Dressler (1993) commented that " Mischobulbon (sic) and Nephelaphyllum resemble the Collabiinae in habit , but their floral features agree with the Bletiinae ". The position in Collabiinae of a large number of the genera that Dressler (1993) placed in Bletiinae (Arthuseae) thus begins to make sense. Vegetatively , these genera form a coherent group, but their floral differences are due to parallelisms with those of other subtribes .

Dressler (1993) placed Arundina and Dilochia together in Arundinae and considered this tribe to be one of the " misfits and leftovers" along with Collabiinae and Pogoniinae . We include Arundina in Arethusinae and Dilochia in Coelogyniae , but we admit that the evidence at this time for a clear demarcation between these two substribes is nuclear (see also Cameron et al., 1999;van den Berg et al., 2000; Goldman et al ., 2001) and may have to be abandoned in favour of Arethuseae s.l.

Our circumscription of Podochileae is also highly at variance with previous circumscriptions. On the basis of Cameron et al. (1999) and van den Berg et al. (2000), we include three subtribes here: Eriiae , Podochilinae and Thelasinae . It is also possible that Bulbophyllinae , Dendrobiinae and Malaxidae fall with this tribe , in which case we would favour including them in Malaxideae (the oldest name). These four groups lack tegular stipes (but some have hamular stipes ), and Malaxideae and Dendrobiinae typically have naked pollinia . All of these formed a clade(with Thelasinae and Ridleyellinae (van den Berg et al. 2000). Eriinae include Pseuderia (Yukawa et al., 1996), and generic delimitaion in the subtribe is clearly problematic and under revision (Ng , 2002).

Epidendreae were studied intensively by van den Berg (2000) , and his results relying upon rbcL, mat K , trnL-F and ITS provide a definitive circumscription . His taxonomic samplying was also extensive and included nearly every New World genus putatively referred to Epidendreae (sensu Dressler , 1993). In these ,Epidendreae are restricted to the New World (Epidendreae I sensu Dressler, 1993) and should probably include both Chysis (Chysinae ; Arethuseae ) and Coelia (Coeliinae ), although at present we cannot say clearly how these two genera fit into the larger picture of the tribe . Several genera within Laeliinae required new circumscriptions (Encyclia , Laelia , and Sophronitis ; Higgins, 1997; van den Berg et al ., 2000), and further work on generic delimitation is still required . Laeliinae should include both Meiracyllium (Meiracylliinae) and Arpophyllum (Arpophyllinae ), but Helleriella p.p., Isochilus, and Ponera (the last perhaps not monophyletic ) should be removed to their own subtribe, Ponerinae . Basiphyllaea (Laeliinae ) also falls outside Laeliinae ), and it ,plus Blettia and Heaxalectris , could be placed in their own subtribe Bletiinae (Bletia as currently circumscribed is possibly not monophyletic ). Pleurothallidinae sensu Dressler (1993) are monophyletic , and their sister clade is composed of Dilomilis and Neocogniauxia (and presumably Tomzanonia as well ; all previously Laeliinae sensu Dressler , 1993). Rather than describe a new subtri composed of such a small number of taxa , we propose to add these three genera to Pleurothallidinae , a circumscription recommended by Pridgeon , Solano and Chase (2001). Dilomilis , Neocogniauxia and Tomzanonia do not have all the synapomorphies of Pleurothallidenae , thu they do share some. Generic limits in Pleurothallidnae also required drastic re-circumscription in several cases (Pridgeon and Chase , 2002). Thus Epidendreae are now composed of just four subtribes ; Bletiinae , Laeliinae , Pleurothallidinae , the first and last rather small and perhaps better combined into one.

Many Old World Epidendreae (Epidendrae II ; Dressler , 1993) would appear to be related to Vandeae. In contrast , Glomera (Glomerinae) falls near Coelogyninae (Cameron et al. 1999; van den Berg et al., 2000; Goldman et al ., 2001). Agrostophyllum was a member of Podochilinae in Dressler (1981) , in which it was unique in having conical rather than spherical stegmata , but he transferred it (1993) to Glomerinae , which all have conical silica bodies (Moller) and Rasmussen , 1984). Earina was a member of Glomerinae in Dressler (1981, 1993) , and it and Afrostophyllum come out together (Freudenstein et al., 2001;van den Berg et al;2003), but neither has anything to do with either Glomera or Podochilinae (we replace the name Glomerinae here with the name Agrostophyllinae because the type genus is not a member of this group). Agrostophyllum and Earina (and we assume the other members of Glomerinae sensu Dressler , 1993).Vegetatively , Earina , Agrostophyllum and Polystachyinae are similar , and they all share elaters (Halle , 1986). Evidence seems to be mounting that Earina and Agrostophyllum (Agrostophyllinae ) are related to the vandoid orchids, which also include Polystachyinae (Fig. 1) . It could turn out that Epidendreae II sensu Dressler (1993) are also monophyletic, in which case Polystachyeae might be the most appropriate name.

Vandoideae sensu Dressler (1981) included Corallorhizinae (the genera of the latter are here included in an expanded Calypsoeae ; see above ), but in nearly all analyses we have recovered a group that consists of Vandeae plus Cymbidieae(including Maxillarieae; see below), which are more or less Vandoideae sensu Dressler (1981) . Dressler soon (1983) repudiated his concept of Vandoideae , and his 1993 classification split these two groups into what he then termed " cymbidioid" and " epidendroid " phylads , which had Cymbidieae and Vandeae as the ulimate members of each emerging from studies of DNA sequences.

The most important character that Dressler relied upon in his delimitation of Vandoideae was the manner in which the anther bacame incumbent (bent down) in development , such that Vandoideae appeared not to be incumbent , whereas Epidendroideae sensu Dressler (1981) were clearly in anther development , so that they appeared not to be incumbent . Moller and Rasmussen (1984) showed that Vandeae shared spherical stegmata (silica bodies) with Dendrobiinae and Eriinae , and this convinced Dressler (1993) that this concept of Vandoideae was polyphyletic , so he proposed instead that advanced Epidendroideae were composed of two " phylads" , " cymbidioid" and " dendrobioid " . Our re-unification of Vandeae with Cymbidieae (in an unnamed group,see below; Fig. 1 ) places together nearly all those taxa with lateral inflorescence , reduced anther partitions , and complex pollinaria composed of viscidium , stipe and two or four superposed pollina. These characters also appear elsewhere (e.g. Calypso and Corallorhizinae sensu Dressler , 1981), but they are nearly universal within Vandoideae. Neverthless , it is likely that the similarly complex within the vandoid complex . The morphological analysis of Freudenstein and Rasmussen (1999) also recovered a Vandeae/ Cymbidieae clade, although they stated that there were insufficient characters in their analysis to make this a reliable result. At present , we have enough evidence to determine that recognition of Vandoideae because it would make Epidendroideae paraphyletic , so we would not propose to reinstate Vandoideae because it would necessitate the description of several other , mostly small subfamilies . It is ironic that Dressler's earlier classification recognised clade later abandoned(1993) that we now recover in the DNA results. Spherical stegmata apparently have evolved at least twice from conical stegmata , and they have been lost in most groups in which a transition to ephemeral stems/leaves has occurred (e.g. Calypsoeae, Gastrodieae and Malaxidae).

Relationships of the subtribes within Vandeae have yet to receive sufficient attention to determine if they are monophyletic ,but they would appear to be (Cameron et al ., 1999, in which taxonomic sampling was the least possible to evaluate subtribal monophyly : Aeridinae , three genera , and Angraecinae and Aerangidinae , two genera each). Generic relationships and delimitation within the subtribes are likewise unexplored in molecular studies , but some work on Vandeae is underway.

In contrast , Cymbidieae are relatively well studied .This group was clearly monophyletic in the rbcL study of Cameron et al. (1999).Cymbidieae sensu Dressler (1993) are paraphyletic to Maxillarieae, and hence we recognise here an expanded Cymbidieae. Likewise Cyrtopodiinae sensu Dressler (1993) are paraphyletic to all the rest ,so we have dismembered them into three tentative subtribes ; Bromheadia is unrevaluated , so we have left it an in Dressler (1993). Whitten scheme here. The only significant differences from that of Dressler (1993) are the recognition of Coeliopsidinae (these genera were included previously in Stanhopeinae ) and submersion of Cryptarrhenianae and Lycastinae im Maxillariinae and Ornithocephalinae , Pachyphyllinae adn Telipogoninae in Oncidiinae . More extensive sampling than in Whitten et al. (2000) for Oncidiinae convincingly demonstrates that Ornithocephalinae , Pachyphyllinae and Telipogoninae are deeply and Telipogoninae are compatible with those found in some genera of Oncidiinae , even though pollinaria with four pollinia in Ornithocephalinae and Telipogoninae caused many previous authors to place them in a tribe with other groups with four pollinia , such as Maxillariinae and Zygopetalinae (Dressler , 1981, for example).

Generic delimitaton in several subtribes has also been studied . Whitten et al. (2000) demonstrated that generic limits in Stanhopeinae accord nearly perfectly with DNA results , as was also ture in the earlier work on Catasetinae (Chase and Hills, 1992; Pridgeon and Chase , 1998), so DNA results do not contradict previous generic schemes based on (intuitively interpreted) morphological information in all cases .Onchidiinae (Williams et al., 2001)are a good example in which many genera have long been thought unsatisfactorily circumscribed (Garay and Stacy, 1976; Chase , 1986; `1987), so the gross polyphyly of the two largest genera , Odontoglossum and Oncidium , came as a surprise to no one .Our list of Oncidiinae genera in the Appendix reflects some of the recent nomenclatural changes , but many more are planned to bring generic delimitation into the line with a strict concept of monophyly . Likewise , many changes are in store for Eulophiinae (Cribb, Pridgeon , Norup and Chase , in prep), Maxillariinae (Whitten, Atwood et al., in prep)and Zygpetalinae (whitten Dressler , Williams et al., in prep.).

3.Conclusions

All of these changes in taxonomy will be reflected in Genera Orchidacearum (Pridgeon et al .,1999 ; 2001; 2003). We expect the classification as outlined here to be ephemeral(hopefully for not longer than the next five years), but it should serve a useful interim purpose of giving other researchers a better place to start than Dressler (1993) , which in spite of its admirable qualities is out of date . Nevertheless , we still recommend that orchid researchers continue to consult his treatment ; it contains a wealth of information and ideas , many of which are still relevant .

Orchids should be one of the premier groups of flowering plants for evolutionary studies , and the massive amout of DNA data now accumulating are revolutionising out ideas about these wonderful plants . Darwin's next book after On the Origin of Species was focused on orchids, and the reasons for this are clear : orchids should be studied more because they opitomise evolution in its most dynamic aspect , the rapid production of an incredibly diverse array of species . The challenge is to understand how this has come out , and so intensive study of this largest angiosperm family is highly appropriate . We hope that this new classification of the family facilitates research on Orchidaceae in the same manner as have Dressler's previous classification (1981; 1993) and that it stimulates an understanding of the urgent need to conserve these evolutionary marvels.

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-, Kita , K . and Handa ,T. (2000).DNA phylogeny and morphological diversification of Australian Dendrobium (Orchidaceae ).pp. 465-471. In Wilson ,K.L .and Morrison ,D.A. eds .Monocots : systematics and evolution . CSIRO Publishing, Collingwood.

Appendix

Subfamilies , tribes ,and subtribes of the Orchidaceae with genera recognised . Genera for which no DNA sequence data are marked with those of which DNA information is available but for which only a tentative placement can be made are placed in (parentheses). Published , in press or submitted studies focusing on some or most of the genera in the various groups are cited by the appropriate tribes and subtribes as well as in the text.

(Unplaced genera are placed following the higher order in which they are thought to be best placed .)Numbers in (brackets) refer tothe number of species listed by Govaerts (2003) with some genera (number in Italics) updated from recent literature . The numbers presented here are likely to be consercative .

Orchidaceae(24, 910)

Apostasioideae(16),Apostasia (7). Neuwiedia(9)

Vanilloideae (249),(Cameron et al., 1999; 2000)

Pogoniieae (77).Clestes (63),Duckeela(3),Isotria (2) ,Pogoniopsis (2) Vanillineae (172),Clematepistephium (1),Crytosia(5),Dictyophyllaria (1),Epistephium (23), Eriaxis (1),Erythrorchis (3),Galeola (7),Lecanorchis (14) ,Pseudovanilla(8), Vanilla (109)

Cypripedioideae (155), (Cox et al., 1999), Cypripedium (50), Mexipedium(1),Paphiopedilum(77), Phragmipedium (21),Selenipedium(5)

Orchidoideae (4, 704)

Cranichideae (1,643), (Salazar et al., 2001, 2003)

Goodyerinae (651), Aenhenrya (1),Anoectochilus (45), Aspidogyne(19), Chamaegastrodia (5), Cheirostylis (47),Cystorchis (21), Danhatchia (1) , Dossinia (1), Erythrodes (88) ,Eurycentrum (7),Evrardianthe (1), gonatostylis (2) , Goodyera (96) ,Gymnochilus (3) ,halleorchis (1),Herpysma (1), Hetaeria (32), Hylophila (10), Kreodanthus (6) ,Kuhlhasseltia (12), Lepidogyne (1),Ligeophila (9) , Ludisia (1) , Macodest (9) ,Microchilus (2) ,Moerenhoutia (13) , Myrmorchis (16) ,Odontochilus (3) , Orchipedum,(2) , Pachyplectron (3), papuaea (1) ,Platylepis (8) ,Platyhelys (10) ,Pristiglottis (22) , Rhmphorhynchus (1) ,Rhomboda (19) , Stephanothelyst (2).Tuberolabium (13) , Vrydagznea (41) ,Zeuxine (76)

Spiranthinae (470) ,Aracamunia (1) , Aulosepalum (6) ,Beloglottis (7) ,Brachystele (20) ,buchtienia (3) ,Coccineorchis (4) Cotylolabium (1) ,Cybebus (1) ,Cyclopogon (82) , Degranvillea (1) , Dieregyne (19) ,Dichromanthus (1) . Discyphaus (1) , Eltroplectris (13) ,Eurystyles (18) , Funkiella (4) ,Hapalorchis (8) ,Helonoma (2) , Kionophyton (4) , Lankesterella (11) , Lyroglossa (2) ,Mesadenella (7) ,Mesadenus (6) , Microthelys (6), Odontorrhynchus (6), Pelexia (77), Physogyne (3) ,Pseudogoodyera (2), Pteroglossa (9),Sacoila (5), Sarcoglottis (41),Sauroglossum (12) ,Schiedeella (21),Skeptrostachys (2) , Thelyschista (1),Veyretia (9) , Wallnoeferia (9) , Wallnoeferia (1)

Manniellinae (1) , Manniella (1)

Cranichidinae (268), Aa (27), Altensteinia (7), Baskervilla (12), coilochilus (1) , Cranichis (54), Exalaria (1), Fuertesiella (1), Gomphichist (25),Myrosmodest (10), nothostele (1) , Ponthieva (57) , Porphyrostachys (2) Prescottia (31),Pseudocentrum (7), pseudocranichist (1) ,Pterichis (20) ,Solenocentrum (4) , Stenoptera (7)

Pterostylidinae (160) , Pterostylis (160)

Chloraeinae (84) ,bipinnula (11), Chloraea (51),gavilea (14), Geoblasta (2),Megastylis (in part )(6)

Galeottiellinae (3) ,Galeottiella(3)

Diuridea (874) ,(Kores et al ., 2001; Clements et al ., 2002)

Acianthinae (170), Acianthus (26) , Corybas (125), Corybas (125), Cyrtostylis (6) , Stigmatodactylus (11), Townsonia (2)

Caladeniinae (281),Adenochilus (2) , Aporostylis (1),Cladenia (250), Cyanicula (10) , Elythranthera (2) , Eriochilus (12), Glossodia (2),Leptoceras (1), Praecoxanthus (1), Cryptostylidinae (26),Coilochilus (1),Cryptochilus (1),Cryptostylis (25)

Thelymitrinate (166) ,Arthrochilus (10) ,Burnettia (1) ,Caleana (1), Calochilus (23), Chiloglottis(27),Drakaea (9),Leporella (1),Lyperanthus (2),Megastylis (in part )(1), Paracaleana (9), Pyrorchis (2), Rimacola (1),Spiculaea (1),Thelymitra(78)

Diuridinae (61) , Diuris (58) , Epiblema (1), Orthoceras (2)

Prasophyllinae (168),Genoplesium(45) ,Microtis (18), Prasophyllum (105)

Rhizanthellinae (2),Rhizanthella (2)

Codonorchideae (2), Codonorchis (2)

Orchideae (2, 185) , (Bateman et al ., 1998; Douzery et al ., 1999 ; Pridgeon et al ., 1998), Ponerorchis (17)

Brownleeinae (83), Brownleea (7) ,Disperis (76)

Disinae (220) ,Ceratandra (6) Corycium (15) ,Disa (169) ,Evotella (1) , Huttonanea (5) , Pterygodium (18) , Schizodium (6)

orchidinae (1,865), Aceratorchis (2) ,Amerorchis (1) , Amitostigma (28) ,Anacamptis (13) ,Androcorys (6) ,Aorchis (2) , Arnottia (4) Barlia (2) , Bartholina (2) ,Benthamia (31) , Bonatea (17) ,Brachycorythis (36),Centrostigma (3),Chamorchis (1) ,Chusua (20),Comperia (1), Cynorkis (158), Dactylorhiza (48),Diphylax (4) , Diplomeris (5), Galearis (1) ,Gennaria (1) , Gymnadenia (24) , Habenaria (848) ,Hemipilia (18)Herminium (28),Himantoglossum (7),Holothrix (44),Megalorchis (1) ,neobolusia (4) , Neotinea (4), Neottianthe (8), Ophrys (60), Orchis (27), Pachites (2), pecteilis (4), Peristylus (102) , Physoceras (11) ,Platanthera (!35),platycoryne (17) ,Porolabium (1),Pseudorchis (1) ,Roeperocharis (5), Satyrium(1),Symphyosephalum (1) ,Thulinia (1) ,Traunsteinera (2), tsaiorchis (1) ,Tylostigma (8)

Epidendroideae (19, 785)Claderia (2)

Neottieae (191), Aphyllorchis (21), Cephalanthera (20),Epipactis (64),Limodorum (3),Neottia (63), [Palmorchis (20)]

Sobralieae (237), Elleanthus (106) . Epilyna (3),Sertifera (8),Sobralia (120)

Tropidieae (35), Corymborkis (6),Tropidia (29)

Triphoreae (28),Diceratostele (1),Monophyllorchis (2), Psilochilus (7) ,Triphora (18)

Nervilieae (73) , Nervilia (65), [Epipogium (4),Silvorchis (1),Stereosandra (1),Xerorchis (2)]

Gastrodieae (70) ,(Molvray et al ., 2000), Auxopus (3) ,Didymoplexiella (6), Didymoplexix (17), Gastrodia (41) Neoclemensia (1) ,Uleiorchis (2)

Calypsoeae (70),(Goldman et al ., 2001),Aplectrum (1),Calypso (1),Changnienia (1), Corallorhiza (11),Cremastra (3) ,Dactylostalix (1) ,Didiciea (2),Ephippianthus (2), Govenia (19), Oreorchis (18),Tipularia (5) ,Yoania (3), [Wullschlaegelia (3)]

Eidendreae (5, 870),(Van den Berg et al., 2000)Chysis (8), Coelia (5)

Ponerinae (22), Helleriella (2), Isochilus (12) , Ponera (8)

Bletiinae (48),Basiphyllaea (7),Bletia (34),Hexalectris (7)

Pleurothallidinae (3, 999), (Pridgeon and Chase , 2001; Pridgeon , Solana and Chase , 2001),Acianthera (131), Anathallis (89), Andinia (24), Anthereon (6),Barbosella (18), Brachionidium (65) ,Chamelophyton (1), Dilomilis (5), Diodonopsis (5), Dracula (111), Dresslerella (9), Dryadella (42),Echinosepala(8) , Frondaria (1) ,Lepanthes (931), Lepanthopsis (38),Masdevallia (507) , Myoxanthus (44), Neocogniauxia (2),Octomeria (143),Phloephila (14), Echinosepala (8) ,Frondaria (1),Lepanthes (931), Lepanthopsis (38),Masdevallia (507),Myoxanthus (44),Neocogniauxia (2),Octomeria (143), Phloeophila (14), Platystele (91),pleurothallis (813) ,Pleruothallopsis (30), Porroglossum (34),Restrepia (48),Restrepiella (1) ,Scaphosalpinx (122) ,Trisetella (22), Zootrophion (12)

Laeliinae (1,788), (van den Berg et al ., 2000),Acrorchis (1),Alamania (1) ,Arpophyllum (4) ,Artorima (1),Barkeria (15),Brassavola (20),Broughtonia (6),Cattleya ( 54), Caularthron (4) ,Dimerandra (6),Dinema (2), Domingoa (2),Encyclia (154), Epidendrum (1,125),Euchile (2), Hagsatera (2), Hexisea (5),Homalopetalum (4), Isabelia (1),Jacquiniella (6) ,Laelia (11),Lanium (9),Leptotes (6) ,Loefgrenianthus (1),Meiracyllium (2), Myrmecophila (10),Nageliella (2) Nanodes (2) Nidema (2) ,Oerstedella (29), Oestlundia (11), Orleanesia (9),Pinelia (4) ,Platyglottis (1), Prosthechea (93),Pseudolaelia (8),Psychilus (15),Pygmaeorchis (2),Quisqueya (4),Renata (1), Rhyncholaelia (2),Scaphyglottis (63),Schomburgkia (15) ,Sophronitis (57), Tetramicra (14)

Podochileae (1,232), (ng ,Yukawa ,etal ., in prep)

Eriinae (725) , Ascidieria (1),Ceratostylis (145),Cryptochilus (4),Epiblastus (22), Eria (404),Mediocalcar (24),Porpax (13),Pseuderia (19),Sarcostoma (5), Stolzia (15).,Trichotosia (73)

Podochilinae (208),Appendicula (142),Chitonochilus (1) ,Poaephyllum (6), Podochilus (59)

Thelasinae (701), (Goldman et al ., 2001)

Arethusinae (9),Anthogonium (1),Arethusa (1) ,Arundina (1),Calopogon (5),Eleorchis (1)

Coelogyninae (692),(Gravendeel et al ., 2001), Bletilla (5) ,Bracisepalum (2),Bulleyia (1),Chelonistele (12),Coelogyne (182),Dendrochilum (264), Dickasonia (1),Dilochia (8), Entomophobia (1) ,Geesinkorchis (2) , Glomera (127), Ischnogyne (1),Nabaluia (3), Neogyna (1),Otochilus (5),Panisea (8),Pholidota (43),Pleione (20), Thunia (5)

Malaxideae (1,158), Crossoglossa (21),Hippeophyllum (13),liparis (418),Malaxis (395), Oberonia (308),Orestias (3)Risleya (1)

Cymbidieae (3, 814),(Whitten et al ., 2000; Pridgeon et al ., in prep)

Bromheadiinae (28)Bromheadia (28)

Catasetinae (367), (Chase and Hills, 1992 ; Pridgeon and Chase , 1998), Catasetum (157),Clowesia (7),Cycnoches (33),Cyrtopodium (44), Dressleria (10), Galeandra (34), Grobya (4) ,Mormodes (78)

Cymbidiinae (67), Cymbidium (51),Grammatophyllum (11),Graphorkis (4),Porphyroglottis (1)

Eulophiinae (316),Acriopsis (7), Acrolophia (7), Ansellia (1),Cymbidiella (3), Cynaeorchis (2) ,Dipodium (24),Eulophia (211),Eulophiella (5),Geodorum (13), Grammangis (2) Oeceoclades (38),Thecopus (2),thecostele (1)

Eriopsidinae (5),Eriopsis (5)

Oncidiinae (1,589), (Sosa et al ., 2001; Williams et al ., 2001; Williams ,Chase and Whitten , 2001), Ada (16),Amparoa (1),Antillanorchis (1), Aspasia (7),brachtia (7), Brassia (34),Caluera (2),Capanemia (14),Caucaea (20),Centroglossa (5), Chytroglossa (3),Cischweinfia (11) ,Cochlioda (7),Comparettia (7),Cuitlauzina (1), Cypholoron (2),Cyrtochiloides (3),Cyrtochilum (119) ,Diadenium (2) ,Dignathe (1),Dunstervillea (1),Eloyella (6),Erycina (7),fernandezia (9),Gomesa (17), Goniochilus (1) ,Hintonella (1),Hofmeisterella (1)Hybochilus (1),Ionopsis (6), Leochilus (11),Lockhartia (27),Macradenia (11),Macroclinium (38), mesospinidium (7) Miltonia (10) ,Miltoniopsis (5),Neokoehleria (10),Notylia (58) ,Odontoglossum (69),Oncidium (336) ,Ornithocephalus (44) ,Osmoglossum (7),Otoglossum(13), Pachyphyllum (39),Palumbina (1),Papperitzia (1) Pfitzeria (1) ,Phymatidium (8) , Platyrhiza (2) ,Plectrophora (10), Polumnina (1) ,Papperitzia (1) , Pfitzeria (1),Phymatidium (8), Platyrhiza (2),Plectrophora (10),Polyotidium (1),Psychopsiella (1),Psychopsis (5), Pterostemma (2),Quekettia (7), Raycadenco (1) ,Rodriguezia (48),Rodrigueziella (6), Rpdrogueziopsis (2),Rossioglossum (4),Rauhiella (3) ,Rhynchostele (16),Sanderella (2),Saundersia (3),Scelochiloidest (3),Scelochilopsis (1), Scelochilus(48), Seegeriella (1),Sigmatorthos (1), Suarezia (1), Sutrina (1),Symphyglossum (4),Systeloglossum (5),Telipogon (133),Thysanoglossa (2),Ticoglossum (2),Tolumnia (36),Trichocentrum (69), Trichoceros (9),Trichopilia (26),Trizeuxis (1),Pityphyllum (5), Rudolfiella (6) ,Scuticaria (8),Teuscheria (7),Trigonidium (14),Xylobium (32)

Stanhopeinae (257),(Whitten et al ., 2000) ,Acineta (15) ,Braemia (1),Cirrhaea (9), Coryanthes (38),Embreea (1),Gongora (58),Horichia (1),Houlletia (9),Kegeliella (3), Lacaena (2),Leuckelia (1),Lueddemannia (1),Paphinia (16),Polucycnis (15),Schlimmia (8),Sievekingia (16),Soterosanthus (1),Paphinia (16) ,Polycycnis (15),Schlimmia (8),Sievekingia (16),Soterosanthus (1),Stanhopea (55),trevoria (6),Vasqueziella (1)

Coeliopsidinae (18) ,(Whitten et al ., 2000),Coeliopsis (1),Lycomormium (6),Peristeria (11)

Zygopetalinae (418),Aganisia (3),Batemannia (5),Benzingia (2)Bollea (12). Chaubardia (5),Chaubardiella (8) ., Cheiradenia (1).Chondrorhyncha (30), Chondroscaphe (12),Cochleanthes (14),Cryptarrhena (4),Dichaea (110),Dodsonia Koellensteinia (19),Neogardneria (19),Stenia (18),Vargasiella (2),Warrea (4),Warreella (2),Warreopsis (4),Zygosepalum (8)

Vandeae (2,341)Christensonia (1)

Polystachyinae (228),Hederorkis (2),Imerinaea (1).Neobenthamia (1),Polystachya (224)

Aeridinae (1,352),Abdominea (1),Acampe (7), Adenoncos (16),Aerides (25), Amesiella (3),Arachnis (11),Armodorum (3),Ascocentrum (13) ,Ascochilopsis (2)Ascochilus (6) ,Ascoglossum (1),Biermannia (10),Bogoria (4),Brachypeza (7),Calymmanthera (5),Ceratocentron (1),Ceratochilus (2),Chameaeanthus (2),Chiloschista (20),Chroniochilus (4),Cleisocentron (3),Cleisomeria (2),(2) Diplocentrum (2) Diploprora (2),Drayadorchis (4),Drymoanthus (4).Dyakia (1), Eparmatostigma (1)Esmeralda (2),Gastrochilus (55),Grosourdya (9),Gunnarella (9),Gunnarella (9),Haraella (1),Holcoglossum (10) ,Hygrochilus (1)Hymenorchis (10) .Lesliea (1),Loxomorchis (3) ,Luisia (39) , Macropodanthus (6) ,Malleola (29) ,megalotus (1) ,Micropera (18) ,Microsaccus (13) ,Microtatorchis (50) ,Mobilabium (1) , Neofinetia (2),Nothodoritis (1),Omoea (2) ,Ornithochilus (3) ,Papilionathe (10) ,Papillilabium (1) ,Paraphalaenopsis (4) ,Parapteroceras (5) ,Pelatantheria (7) ,Pennilabium (10),Peristeranthus (1) ,Phalaenopsis (62) ,Phragmorchis (1) ,Schoenorchis (26) ,Sedirea (2) ,Seidenfadenia (1) ,Smithsonia (3) ,Smitinandia (3) ,Staurochilus (14) ,Stereochilus (7) ,Taeniophyllum (185) ,Thrixspermum (144) , Trichoglottis (64) ,Tuberolabium (12) ,Uncifera (6) ,Vanda (57), Vandopsis (5) , Ventricularia (2),Xenikophyton (1)

Angraecinae (445) ,Aeranthes (47) , Ambrella (1) ,Angraecum (219) ,Bonniera (2) , Clayptrochilum (2) ,Campylocentrum (73) ,Cryptopus (4) ,Dendrophylax (9) , harrisella (3) ,Jumellea (58) ,Lemurella (4) ,lemurorchis (1) .Listrostachys (2)., Neobathiea (5) Oeonia (6) Oeoniella (2) Ossiculum (1) ,Sobennikoffia (4)

Aerangidinae (315) ,Aerangis (49 ) , Ancistrorhynchus (16) ,Angraecopsis (21) , Beclardia (1) Bolusiella (6) ,Cardichilos (1) Chamaeangist (10) Chauliodont (1) , Cribbia (4) ,Cyrtorchis (15) Diaphananthe ((24) ,Dinklageella (3) ,Distylodont (1) Eggelingia (3) ,Eurychone (2) ,margelliantha (5),Microcoelia (29),Microterangis (7),Rhaesteria (1) ,Rhipidoglossum (37) ,Solenangis (6) ,Sphyrarhynchus (1) , Summerhayesia (2) ,Taeniorrhiza (1) ,Triceratorhynchus (1) ,Tridactyle (43), Ypsilopus (5)

Unplaced subtribes within Epidendroideae (3, 963)

Agrostophyllinae (196) ,Adrorhizon (1), Aglossorhyncha (13),Agrostophyllum (91), Earina (6).Glossorhyncha (80) ,Ischocentrum (2),Sepalosiphont (1), Sirhookera (2) Dendrobiinae (3,332),(Yukawa et al ., 2000)Bulbophyllum (1,789),Cadetia (60), Chaseella (1) ,Dactylorhynchus (1),Dendrobium (1, 184), Diplocaulobium (99), Drymoda (3),Epigeneium (38),Flickingeria (69),Genyorchis (7),Jejosephia (1), Monomera (3)Monosepalum (3) ,Pedilochilus (35),Saccoglossum (5), Sunipia (22) Triast (12)

Collabiinae (435), (Goldman et al., 2001),Acanthephippium (12) ,Ancistrochilus (2), Aulostylis (11),Calanthe (187) ,Cephalantheropsis (5),Chrysoglossum (4),Collabium (14).Diglyphosa (3),Eriodes (1),Gastrorchis (9),Hancokia (1) .Ipsea (3) , Mischobulbum (9),Nephelaphyllum (12), Pachystoma (1),Phaius (48), Plocoglottis (39) ,Spathoglottis (45) ,Tainia (29).