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Assembling the Tree of Life: Orchidaceae Orchid Tree:
    a phylogeny of epiphytes (mostly) on the tree of life
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Management Plan

Assembling the Tree of Life for Orchidaceae is a large-scale, cooperative project among three US institutions (Univ. of Florida, Ohio State Univ., and New York Botanical Garden) in collaboration with investigators in the United Kingdom, Australia, Brazil, Costa Rica, Denmark, Germany, Japan, and Mexico. Together, this group of researchers have a long and productive history of collaborative research projects on various aspects of the biology of Orchidaceae. The main PIs and collaborators involve most workers in orchid molecular systematics today in Europe, UK, USA, Mexico, Central America, and Brazil. The group of investigators has been assembled to take advantage of their diversity of backgrounds, experience and expertise, and resources already assembled. Additional collaborators are welcomed. Orchidaceae is an ideal group of organisms around which to assemble educational efforts concerning the importance of evolution, conservation, systematics, phylogenetics, and biodiversity, as well as to involve students and investigators from tropical countries.

1. Resources

1. A. Major core personnel: The major core personnel at USA and UK institutions range from junior to senior faculty members who have diverse backgrounds at major research and educational institutions from public universities to two of the largest botanical gardens in the world.

Norris H. Williams will serve as lead PI on the project. He has an extensive background in orchid biology, including orchid anatomy, palynology, pollination biology, chemical ecology, and molecular systematics. He served for nine years as Chair of the Research Department of the Florida Museum of Natural History and has administrative experience with large, complex groups. He has developed extensive contacts throughout Central and South America during the past 40 years, has recruited students from Mexico to Brazil, and has active collaborations in Mexico, Costa Rica, Panama, Ecuador, Peru, and Brazil. He will oversee organization and administration of the project and will supervise database and website organization and development. He will be responsible for recruiting minority graduate students from two of the historically black colleges and universities in Florida (Florida A&M University and Bethune-Cookman College).

W. Mark Whitten has extensive field and lab experience in orchid systematics, pollination biology, orchid chemical ecology, and molecular systematics. He will supervise DNA aliquot distribution, voucher databasing, PCR and sequencing troubleshooting, GenBank data entry, and matrix construction. He and Williams will be responsible for ca. 1/3 of sequence data collection.

John V. Freudenstein has extensive experience in morphological and molecular systematics, data analysis, and methods of phylogenetic analysis. He will lead the morphological data collection and analyses and will be responsible for ca. 1/3 of sequence data collection. He will also lead the data analysis at OSU.

Mario Lauria is a computer scientist who has extensive experience that covers different aspects of super computing systems, PC cluster technology and architecture, system software, applications - and the intersection of cluster architecture and Grid computing. He has been developing new ways of building and programming clusters specifically to solve demanding computational biology problems including sequence alignment. He will develop the software tools for the project and will work with PI Freudenstein to conduct the major data analyses.

Kenneth M. Cameron has extensive experience with higher level phylogenetics of Orchidaceae and has extensive field experience in Asia and Australia. He will oversee sequence data collection for ca. 1/3 of taxa and will be responsible for education and outreach products.

1. B. Collaborators: We have assembled a diverse group of international collaborators who have tremendous experience in various aspects of orchid biology, molecular systematics, and herbarium and field work with Orchidaceae. Letters of collaboration are attached.

Mark W. Chase is Head of the Molecular Systematics Section, Jodrell Laboratory, Royal Botanic Gardens, Kew and is the leader of many major collaborations in angiosperm molecular systematics. He was instrumental in developing large-scale collaborative phylogenetic projects and has extensive experience in orchid systematics. Most orchid molecular phylogenetic workers have collaborated with Mark, and his lab at Kew has played a leading role in training orchid biologists and in assembling the DNA bank critical to success of this project. He and collaborators at Kew (Michael Fay, Phillip Cribb, Dave Roberts, Vincent Savolainen) will consult on taxon sampling, phylogenetic analysis, and studies of biogeography, phylogeography, and molecular evolution.

Richard Bateman (United Kingdom) is a leading authority and researcher on European orchids with an extensive collection of DNA samples and morphological data for these taxa.

Cassio van den Berg (Brazil) has worked extensively on molecular and morphological systematics of the large and horticulturally important Laeliinae and will contribute his expertise and samples.

Germán Carnevali (Mexico) is collaborating with us on studies of Maxillariinae and Laeliinae; he has broad expertise in orchids of Mexico and Venezuela.

Mark A. Clements (Australia) is working on the large and difficult subtribes Diuridae and Dendrobiinae, and has assembled an extensive DNA bank of Australasian species.

Phillip Cribb (UK) is Curator of the Orchid Herbarium at Royal Botanic Gardens, Kew, and is an authority on Cypripedioideae and many African and Asian groups. His cooperation and guidance through the herbarium and library resources at Kew are essential.

Robert Dressler (US, Costa Rica) authored two of the most influential books on orchid phylogeny and evolution and publishes frequently on orchids of Mesoamerica. He is a research associate at UF, and will assist with morphological character evaluation, choice and scoring, and with verification of voucher determinations.

Guenter Gerlach (Germany) has extensive experience with Stanhopeinae and Zygopetalinae and has made (and will make) available critical material from the Munich Botanical Garden.

Barbara Gravendeel (Netherlands) has published on Coelogyninae and is actively working on the large subtribe Bulbophyllinae (over 2000 species).

Eric Hágsater (Mexico) is the authority on Epidendrum (1100+ sp), and has extensive living, herbarium, and DNA accessions of this group and of additional Mexican taxa.

Alexander Kocyan, also at Munich Botanical Garden, has extensive experience in southeast Asian groups and is actively working on the Vandeae.

Gerardo Salazar (Mexico) is essential for his expertise on Mesoamerican taxa, as well as for his expertise on the Cranichidinae.

Tomohisa Yukawa (Japan) Tsukuba Botanical Garden National Science Museum, Tsukuba, is a valuable collaborator with extensive knowledge of East Asian taxa.

Finn Rasmussen (Denmark) has published extensively on orchid morphology and anatomy, and will work closely with John Freudenstein to develop morphological matrices.

Franco Pupulin (Costa Rica) contributes extensive knowledge on systematics and especially of morphology of Central American taxa.

Gustavo Romero (USA) is curator of the Ames orchid herbarium at Harvard, with its excellent collection of types and rich library. Gustavo has broad expertise in neotropical taxa.

Samantha Koehler (Brazil) is actively working on molecular systematics and pollination biology of Brazilian genera. Together, Koehler, Singer, and van den Berg have access to large living collections, DNA banks, and herbarium resources within Brazil.

Jason Keel is currently webmaster for Anthropology at UF. He has long interest and expertise in web and database design, and will supervise and advise on this portion of the project.

Together, the group represents a diverse, world-wide assembly of some of the most active and productive researchers on Orchidaceae.

1. C. Collection resources: A project of this magnitude requires vast collection resources of both herbarium and DNA materials. No field collecting will be required since an enormous collection of vouchered orchid DNA has already been amassed by the PIs and their international collaborators. These specimens and their banked DNAs were collected in compliance with national and international regulations such as CITES; the transfer of DNA samples among institutions will comply with CITES and with Materials Transfer Agreements (MTAs) regulating any specimens. The herbarium collections available at the CITES-registered institutions involved encompass the vast majority of the species and genera to be sampled. The Kew Orchid Checklist (http://www.kew.org/wcsp/home.do) of all currently recognized genera and species [Royal Botanic Gardens, Kew (2004)] is used as our reference. The current link is to the entire live, database-driven monocot checklist. This is a complete list of all described species and genera with included synonymy and forms the basis of our sampling strategy.

1. D. Laboratory facilities: Although some sequencing will be done at UF, NYBG, and OSU, all of which have well-equipped labs and sequencing facilities, in order to reduce costs, we will outsource the majority of the sequencing to the University of Washington High Throughput Genomics Unit (see budget for quote and details). Details of facilities are given in the appropriate place in the proposal.

1. E. Computational facilities: Because of the size of the data matrix, we will use both our individual computers and clusters for initial analyses. Additional facilities include the clusters and supercomputer facility at OSU which will be used for the large, combined analyses (see Facilities). Computation times are actually often decreased by the use of additional genes, as multi-gene combined analyses have been shown to run faster than single gene data sets for large taxon data sets.

2. Database: Collaborator Jason Keel will ensure compliance with Taxonomic Databases Working Group standards (e.g., Structure of Descriptive Data (SDD), Darwin Core, and Access to Biological Collections Data (ABCD)), as well as standards and specifications that affect the broader community, e.g., eXtensible Markup Language (XML), and OpenGIS Consortium specifications, (Geographic Markup Language (GML) and Web Mapping Service (WMS)). The database will include links to original descriptions, illustrations, photographs, scanned images, morphological characters and states, herbarium data, etc. and will be made available on our web site as the project is in progress in order that all information will be available to collaborators for comment and any necessary corrections. We will have a dedicated server for the project and all sequences, both as raw data and as edited sequences and as incorporated into data matrices, will be posted on a weekly basis to the collaborator web site. Daily backups will be made to an external set of hard drives, as well as to the University of Florida central computer back-up facility. All data remain the property of the researchers until they are satisfied with the quality and editing of the data. Once the researchers are satisfied that the data can be released, sequence data will be sent to GenBank on a monthly basis. Explicit voucher information (collector number, sheet number, and herbarium) will accompany all GenBank entries. As all of the researchers are using Sequencher (GeneCodes), it will be easy to exchange and examine data in contig form among institutions. Images in the database (herbarium specimens, diagnostic line drawings, photographs) will be stored and served over the internet as high resolution zoomable images using JPEG 2000 formats (please see images on our herbarium website http://www.flmnh.ufl.edu/herbarium/cat/digitalimagingprojects.htm). We will purchase dedicated servers for serving these images. The database data and images will be presented in the form of a hyperbolic tree (see below).

3. Website: This project will generate a large (2000 taxon), complex phylogenetic tree with associated images and data. Presenting this tree to other researchers and the public is difficult due to its sheer size; even printing the tree is difficult. The best solution we have found is the hyperbolic tree (not supertree) concept as implemented in the StarTree software from Inxight, which we have already purchased. (See demo at http://www.flmnh.ufl.edu/orchidatol/phylogenetics/plugins.htm). (Please check it out - use your cursor to grab a node and drag it to expand it.) This software allows trees to be presented in a zoomable and searchable format that allows users to easily navigate and manipulate large trees. The nodes and terminals will eventually include images and links to databases. Unlike a static journal publication, this tree will be easily revised when new taxa and new data are added. We feel this method of data presentation is superior to those currently utilized for displaying large phylogenies and is much more easily edited and revised. In addition to our own website, data and resulting trees will be deposited in TreeBase at (http://www.treebase.org/treebase/ and on the Tree of Life web site at http://tolweb.org/tree/phylogeny.html..

The IT technician will work to develop the StarTree Orchid phylogeny and to integrate it with linked databases and will keep the web site up to date and make sure the links all work.

Table 1. ASSIGNMENT OF TAXA (approximately 667 species per lab)

University of Florida
CYMBIDIEAE (3814) included subtribes listed below

Ohio State University
EPIDENDREAE in part (1858)

New York Botanical Garden
EPIDENDREAE in part (1858)

4. Tasks for each team member and time scheduling



Year 2

Year 3

Year 4

Year 5


Recruit minority grad students;begin databasing and web improvements

Matrix construction; analysis; imaging

Coordinate publications for Monocots IV symposium. postdoc

Matrix construction; analysis; imaging; search,

Coordinate publications, symposia, website


Exchange DNAs; Sequencing & databasing

Sequencing & databasing

Sequencing & databasing

Sequencing & databasing

Publishing; outreach materials



Sequencing & databasing; search postdoc

Sequencing & databasing

Sequencing & databasing

Publishing; outreach materials

NYBG postdoc

(yrs 3-4)


Develop education & outreach materials

Develop education & outreach materials



Sequencing; morph. char. selection; search postdoc

Sequencing; morphological data coll.

Sequencing; morphological data coll.

Sequencing; morphological data analysis

Publishing; outreach materials

OSU postdoc

(yrs 2-3)


Morphological data collection

Morphological data collection & analysis


Lauria (OSU)

Prototype of parallel workflow execution system

add remote storage access

Interface to computational grids using Web Service interfaces

comparative analysis and performance assessment

Final release of software and preparation of documentation

Bioinformatics Tech (1/2 time in last four years)

Construct database & merge with StarTree software

Database implementation & refining

Database implementation & refining

Database implementation & refining

Database finalization; web outreach

You can find a detailed list of genera and number of species to be sequenced, as well as a table of the morphological characters to be used, towards the end of the Management Plan. You will need to scroll towards the bottom of that page to get to the lists.

5. Communication among team members

We plan to give regular presentations at national and international scientific meetings at all stages of the project. These meetings will include the annual Botanical Society of America meetings, the International Monocot Symposium ("Monocots IV" in Copenhagen, 2008), and the World Orchid Conference in Miami in 2008. To minimize travel costs, we will plan annual coordination meetings in conjunction with the annual BSA meetings.

6. Annual milestones for progress

Year 1: Recruit grad students, bioinformatics tech, postdoc search; organize DNAs; begin sequencing 8 regions/lab; initiate independent and combined molecular data matrices;

Year 2: sequence 167 taxa/8 regions/lab; complete database design and implementation;

Year 3: sequence 167taxa/8 regions/lab; focus on morphological matrix;

Year 4: sequence 167taxa/8 regions/lab; assemble final molecular matrix; complete morphological matrix; begin education/outreach products;

Year 5: Final sequencing, final analyses, publications; symposium; education/outreach module distribution; website completion.

7. Curatorial, sequencing, computational and informatic facilities

All three lead institutions have molecular systematics laboratories equipped for routine PCR. All three labs have dedicated computers and software (Sequencher, PAUP*, MacClade, Nona, MrBayes, Lucid, etc.) for sequence acquisition, editing, data matrix construction, and routine analyses. Funds are requested for purchasing two dedicated servers (one for development and one for public serving, plus associated software) for serving hyperbolic trees linked with zoomable high-resolution images and linked databases resulting from the grant work. These servers have a three year warranty and an expected life span of four to five years; they will be replaced in year four to insure physical maintenance of the database server beyond the termination of the project. The cluster and supercomputer facilities at OSU are described in the Lauria Facilities Description.

8. Coordination with foreign-based projects on same or related organisms: Collectively, the PIs and collaborators are working with or are communicating with nearly all workers in orchid phylogenetics. We are actively seeking additional collaborators, especially those from Asia and Africa, who will be encouraged to participate in the project.

9. Training and outreach: Educational and outreach materials will be developed primarily by Cameron and his postdoc at NY, but the Florida Museum of Natural History (co-PIs Williams & Whitten) has a large exhibit program and exhibit hall (http://www.flmnh.ufl.edu/exhibits/); its staff has a successful track record of grant funding for exhibit design and construction, education, and outreach through traveling exhibits. When our ATOL grant is funded, we will pursue separate grant funding for traveling exhibit design and construction in conjunction with the exhibit/education staff at FLMNH and NYBG. In addition, PI Williams will make a concerted effort to recruit minority graduate students from two of the historically black colleges and universities in Florida (Florida A&M University and Bethune-Cookman College). See budget justification for elaboration on this aspect and letters of support.

10. Relation to currently funded projects: Cameron was funded by NSF for work on the vanilloid orchids. That project contributed much of the vital DNA's for this proposal. Williams and Whitten are currently funded (through December 2006) by NSF for work on the large Neotropical Maxillariinae. That project also contributed a large amount of the DNA's needed for this proposal. The Maxillariinae project and this proposal complement each, with this proposal providing greater detail and support for the data generated by the Maxillariinae project and the Maxillariinae project providing DNA's and a framework for this project. Freudenstein is funded for work on Corallorhizinae; this project will also provide greater context for that work.

This ends what is in the actual grant proposal. The table of morphological characters and the table of taxa follow.

Table of morphological characters and character states to be used as a starting point.
The list of taxa to be sampled follows this table.

This table is taken from the paper by Freudenstein & Rasmussen and the comments refer to taxa and characters as studied in that paper.

Freudenstein, J. V. & F. N. Rasmussen. 1999. What does morphology tell us about orchid relationships? -- A cladistic analysis. American Journal of Botany 86: 225-248.

This list of characters represents the beginning of a morphological data set for all Orchidaceae. There are 71 character states organized into 53 characters.

0= root tubers 0=absent 1=present

Root tubers are thickened roots that serve as organs of perennation, producing shoots in subsequent seasons. These structures occur primarily in Diseae, Orchideae, and Diurideae and have been studied intensively (e.g., Irmisch, 1850; Germain de St.-Piérre, 1855; Prillieux, 1865; White, 1907; Ogura, 1953; Pridgeon & Chase, 1995). Triphora was also scored as having a true root tuber, but Pogonia and Isotria were not, even though their roots (unthickened) are known to produce shoots (Ames, 1922). Root thickenings are known from Mesadenella (Pabst & Garay, 1952), Apostasia (Stern & Warcup, 1994), Wullschlaegelia (Freudenstein, unpubl.), and possibly others, but these evidently do not produce new shoots.

1= roots 0=rhizodermis 1=velamen

2= exodermis 0=unthick 1=unif. thick 2=outer thick

3= exodermal cells 0=±isodiametric 1=elongate

4= velamen cell thickenings 0=absent 1=linear 2=circular

These characters derive primarily from the work of Porembski & Barthlott (1988). Instead of coding their ten velamen types as states for the terminals, which would necessarily have to be coded as unordered states of a single character, we chose to code these component features of the variation that they describe.

5= spiranthosomes 0=absent 1=present

This character was first described by Stern et al. (1993a). Spiranthosomes are specialized amyloplasts known only from the spiranthoid orchids. Most of the occurrences of spiranthosomes are taken from Stern et al. (1993a, 1993c) Many of the coded absences, outside of spiranthoid orchids, are assumed, since they have not been described before from other groups.

6= growth pattern 0=sympodial 1=monopodial

Most orchids follow the sympodial growth pattern common in monocots (Holttum, 1955). Notable exceptions to this are Vanilla, which is a monopodial vine with elongate internodes, and members of the Vandeae, which have monopodial growth, but shortened internodes.

7= thickened stem 0=absent 1=present

Stem thickening may be in the form of swelling in aerial stems, as in some species of Dendrobium, as corms (e.g., Aplectrum), or as pseudobulbs. If the stem is notably but uniformly thickened, as in Vanilla, the taxon is not scored as having a thickened stem.

8= no. of thickened internodes 0=several 1=one

This character codes variation in numbers of internodes that comprise a pseudobulb. In many taxa, it is clearly one internode (e.g., Bulbophyllum), while in others (e.g., Catasetum), it can be several. Those taxa with no thickened stem are coded as unknown.

9= phyllotaxy 0=spiral 1=distichous

Phyllotaxy in orchids is described as either spiral or distichous, with the latter supposed to be the more advanced state (Dressler & Dodosn, 1960; Dressler, 1993). We found that in all species examined, the condition at the base of a stem, represented by the first bracts surrounding a bud, is distichous. In some taxa (e.g., Apostasia, Neuwiedia, many spiranthoids and diurids), the phyllotaxy soon shifts to more or less spiral, so that most of the leaves appear spirally arranged. In many of these, the arrangement is not strictly spiral, as was shown by Fuchs and Ziegenspeck (e.g., 1927a, b, c). In other taxa (e.g., species examined of Dactylorhiza, Orchis, many epidendroids), the majority of leaves show a distichous arrangement. Even in those species with distichous leaves, the inflorescence is generally spiral in floral arrangement. In some cases, even the infloresence is distichous (Tropidia effusa Rchb. f., Liparis gibbosa, Phalaenopsis cornu-cervi). Hence the basic model for orchid phyllotaxy appears to be a distichous basal condition on the shoot, with a possible shift to a spiral or pseudo-spiral condition at some point (before inflorescence or not). We scored states according to whether a taxon had spiral (or pseudo-spiral) or distichous phyllotaxy on the leaf-bearing portion of the shoot.

10= leaf morphology 0=flat, "herbaceous" 1=plicate 2= conduplicate

Variation among orchid leaves is more complex than it appears at first glance. Phyllotaxy plays a part in determining mature morphology, and since this was scored as a separate character, it is important not to duplicate this information. Leaves with convolute vernation usually are spirally arranged, while conduplicate leaves have duplicate vernation. The state found in many of the terrestrial spiranthoids and orchidoids has been termed by Dressler (1990c) "nonplicate herbaceous". It describes those leaves that are essentially flat (not conduplicate), without prominent midrib, usually fleshy in texture, and that are the result of convolute vernation. Plicate leaves may result from either convolute or duplicate vernation, and are characterized by the accordion-like pleating of the lamina. Conduplicate leaves, as coded here, are those with a single fold of the lamina at the midrib, and without any plication. There are some conduplicate plicate leaves (e.g., Sobralia), but, again, in order to avoid redundancy, these were simply scored as plicate. The states are unordered.

11= winter leaf 0=absent 1=present

Some genera of terrestrial orchids have a leaf that is produced at the end of the growing season, overwinters, is photosynthetically active when conditions are favorable, and withers at or before flowering. These are primarily members of Corallorhizinae (e.g., Aplectrum, Calypso; cf. Freudenstein, 1994), but occasionally other species show a similar modification (some Spiranthes have overwintering leaves [Rasmussen, unplubl.]).

12= leaf articulation 0=absent 1=present

This character has been discussed and the state noted for each orchid group by Dressler (1981) and reflects the presence or absence of an abscission zone at the base of a leaf. As a general rule, most epiphytic orchids are articulate, while terrestrial species are not.

13= stegmata 0=conical 1=spherical 2=absent

Most of the information on this character is from Møller & Rasmussen (1984). Additional scoring was derived from sources quoted in Solereder and Meyer (1930) and from Stern et al. (1993) for Spiranthoideae. The character seems to be largely uniform within genera, although Dressler & Cook (1988) found conical stegmata in Eria javanica, while other members of the genus are known to have spherical stegmata. Since the outgroup (Hypoxis) does not have stegmata present, the plesiomorphic state in the analysis is absence, although Apostasia and Neuwiedia, considered to be a basal lineage, do have conical stegmata. Because stegmata are always found in association with fibrous support tissue, absence may be due either to their loss of the stegmata themselves (where sclerenchyma is present), or to the loss of sclerenchyma. Since presence of fibrous support tissue in the leaf is scored as a separate character, those taxa that have no leaf sclerenchyma were scored as unknown for the presence of stegmata.

14= leaf fiber bundles 0=present 1=absent

Many orchid leaves have sclerenchyma associated with the vascular bundles, as well as independent fiber bundles, while other leaves have no scelerenchyma. Stern et al. (1993c) used this character in their analysis of Spiranthoideae. Presence of sclerenchyma is often, but not exclusively, associated with large leaf size.

15= leaf abaxial epidermal cells 0=straight 1=wavy

Lavarack (1971) utilized this character in his phenetic study of Australian Orchidaceae (Diurideae), and Stern et al. (1993c) surveyed it in the Spiranthoideae. The states are usually quite distinct, with either clearly sinuous anticlinal walls when viewed in paradermal section, or with straight-polygonal walls (cf. Dressler, 1993, fig. 2-6). Adaxial surface cells may also exhibit the condition, but it is most pronounced on the abaxial surface.

16= subsidiary cells 0=present 1=not distinguishable

Some detailed studies of subsidiary cells in orchids have been done (Williams, 1975, 1979; Rasmussen, 1981, 1987), but not on a broad enough scale to make it possible to score the different developmental patterns for many of our terminals. Hence, we have simply scored the presence/absence of subsidiary cells that are morphologically distinct from surrounding epidermal cells, without regard to their ontogeny.

17= inflorescence position 0=terminal 1=lateral

In most orchids this appears to be a relatively straightforward character, although, as shown by Andersen et al. (1988) for Eria, there may be variation within a genus and the true state can be difficult to ascertain.

18= floral abscission 0=absent 1=present 2=present, stalked

Many orchids have an abscission layer at the base of the pedicel; if the flower is not pollinated, it falls from the inflorescence. As opposed to perianth abscission, floral abscission seems to occur only in a more "advanced" group, the epidendroids. Within the Pleurothallidinae there is a further specialization, in that the abscission zone is located between the pedicel and ovary, so that after the flower falls a distinct stalk remains (Dressler, 1993; Neyland & Urbatsch, 1993). This character was coded as ordered (equivalent to coding the stalked ovary as a distinct character).

19= perianth abscission 0=present 1=absent

As with leaves, the perianth may dehisce from the flower at the summit of the ovary. Dressler (1983) described this feature and noted its occurrence in the "basal" orchid groups.

20= calyculus 0=absent 1=Vanilloid 2=Polystachyoid

The calyculus is a series of small bractlike structures outside of the normal perianth. Because of its resemblance to a small perianth whorl, significant evolutionary implications sometimes have been ascribed to the structure -- such as Lindley's (1847) homologizing the calyculus with an additional floral whorl and consequent reinterpretation of the inner whorls. Soon thereafter, however, Crüger (1849) concluded that the calyculus is not an additional floral whorl. Additionally, Kurzweil (1987a) examined the development of these structures in Neobenthamia, and found no relation to the perianth whorls.

In this study two types of calyculus are recognized. Epistephium and Lecanorchis, and to a lesser extent, Vanilla, have a distinct collar below the perianth (Hashimoto, 1990; Dressler, 1993). The type known from Neobenthamia, and also observed here (and by Kurzweil, 1987a) in Polystachya, is different, being essentially a series of swellings on the ovary valves. A bractlike calyculus has also been reported from some species of Bulbophyllum (e.g., Lindley, 1838; Seidenfaden, 1979; Dressler, 1981).

21= lip slipper-shaped 0=absent 1=present

This distinctive labellum form has the middle portion greatly expanded and the apex and distal margin pulled back toward the column, forming a hollow, shoelike structure. This morphology is characteristic of the Cypripedioideae. Superficially similar structures occur occasionally in the Epidendroideae ( e.g., Calypso), but do not include the inrolled margin that is present in Cypripedioids.

22= apiculate perianth 0=absent 1=present

The perianth apices of Apostasia and Neuwiedia are prolonged into distinctive tips, which are formed by the midrib. To our knowledge this feature has not been discussed before in a phylogenetic context, although it is visible in illustrations of the flowers (de Vogel, 1969), but it seems consistent within these genera.

23= petals carinate 0=present 1=absent

Rasmussen (1982; fig. 61d) illustrated an unusual feature in Vanilla, interlocking sepals and petals. This creates a keel on the adaxial surface of the petal along the midrib that is found in several putatively basal orchid groups.

24= lip-column marginal adnation 0=absent 1=present

In some vanilloid orchids the labellum is fused to the column marginally to varying degree. Other taxa sometimes have labellum column fusion of other types, but this is not included here.

25= dorsal median anther 0=present 1=absent

26= lateral inner anthers 0=present 1=absent

The homology of the functional anthers present in orchids was worked out by Brown (1833), Darwin (1862), Swamy (1948), and Rao (1974). Further investigation was not undertaken here; we assume that all monandrous orchids have A1, all diandrous orchids have a1 and a2, and triandrous orchids have A1, a1, and a2. No evidence has ever been presented to substantiate Garay's (1960) claim that two lateral anthers of the outer whorl are present in Satyrium. Occasional unusual situations, such as the presence of a third functional anther in Phragmipedium lindenii and the putatively peloric forms observed by Chen (1982) were not included.

27= anther orientation 0=erect 1=bend late 2=bend early

Anther bending (incumbency) during ontogeny has been discussed by Hirmer (1920) and Dressler (1981, 1986b, 1993). Although there has been some question of whether "advanced" epidendroids bend at all (Dressler, 1981), further study suggests that these taxa bend very early (Dressler, 1986; Kurzweil, 1987a). We distinguish between early and late bending following Kurzweil's (1987a:438) stage 2-3 distinction, relative to the time of column elongation. This character is ordered based on ontogenetic information -- the erect state is more general than the incumbent state (cf. Kurzweil, 1987a).

28= operculate anther 0=absent 1=present

Typical lilioid anthers dehisce by slits, releasing pollen without also shedding parts of the anther wall. Some orchid anthers (Apostasioideae, Cypripedioideae, Orchidoideae, Spiranthoideae - N.B. this paper was written before we knew that the Spiranthoideae was included in the Orchidoideae and before Vanilloideae was recognized as a subfamily) also have basically this type of dehiscence. Incumbent anthers usually need to be physically disturbed to allow the pollinia to be removed. In some taxa with incumbent anthers, the anther develops in tight proximity to the clinandrium; in order for pollinia to be released, the anther (sometimes called the "anther cap") must be removed. This removable type of anther is called an operculate anther.

29= Endothecial thickenings 1 0=other 1=int 2=type II

30= Endothecial thickenings 2 0=other 1=type III/IV

Endothecial thickening morphology in orchids was surveyed by Freudenstein (1991). Here the distinctive type II thickening is coded as one character, and the types III/IV and their intermediates are coded as a second character. State 1 in character 29 is for those thickenings that are intermediate in morphology between types I and II. Because it is a eesentially a morphocline, the character is coded as ordered.

31= basal caudicles 0=absent 1=present

Caudicles are pollinium stalks that are composed of pollen and/or pollen-derived substances, as opposed to rostellar tissue (Richard, 1817; Rasmussen, 1986). In epidendroid and spiranthoid orchids they are produced apically in the anther, due to the bending of the anther or the apical position of the rostellum, respectively. In orchidoids the caudicles are basal extensions of the pollinia.

32= hammer stipe 0=absent 1=present

A distinctive stipe with a "hammer"-like morphology was identified by Rasmussen (1986) in Sunipia. We found this type also in Genyorchis.

33= epidermal stipe/tegula 0=absent 1=present

Rasmussen (1986) has distinguished between the tegula, a pollinium strap formed from the abaxial cuticle of the rostellum, and a hamulus, the apex of the rostellum itself. Rasmussen and Freudenstein (unpubl.) have since distinguished other varieties of pollinium strap. One that is common in epidendroids is an epidermal strap that consists of a varying number of epidermally-derived cell layers plus the cuticle. Because it is sometimes difficult to distinguish between this type and a true tegula, we have combined the two in this character.

34= pollen unit 0=monad 1=tetrad

Mature pollen unit was described by Schill & Pfeiffer (1977) for a large number of species; others were reported in Newton & Williams (1978), Ackerman & Williams (1980, 1981), and Hesse et al., (1989). Ackerman & Williams (1981) described cases of some diurids (e.g., Caladenia) in which both states occur. Wolter & Schill (1986) suggested that the occurrence of tetrads in orchid pollen may be due to paedomorphosis.

35= pollen tectum 0=reticulate 1=smooth

Pollen structure has been described by Schill & Pfeiffer (1977), Burns-Balogh (1983), Hesse et al. (1989) and Zavada (1990). Rather than focus on details of the tectal structure, which has not been studied in enough genera, we simply scored the appearance of the pollen grains, whether reticulate or smooth.

36= pollen apertures 0=colpate/sulcate 1=porate 2=inaperturate 3=polyporate

The aperture state of orchid pollen was taken from Erdtman (1944, 1966), Schill & Pfeiffer (1977), Newton & Williams (1978), Ackerman & Williams (1980), and Burns-Balogh et al. (1987). The greater proportion of orchid pollen is inaperturate (Schill & Pfeiffer, 1977), but putatively basal groups have sulcate-colpate or porate pollen (Newton & Williams, 1978; Burns-Balogh et al., 1987). Some of the vanilloids (Vanilla, Epistephium, Lecanorchis) have been reported to have polyaperturate pollen (Erdtman, 1944, 1966; Schill & Pfeiffer, 1977; Ackerman & Williams, 1980), a feature otherwise unknown among orchids.

37= operculate colpus 0=absent 1=present

Burns-Balogh & Funk (1986) utilized this character in their analysis. This pollen feature was described and illustrated by Schill (1978) and Newton & Williams (1978) and only appears in Apostasia and Neuwiedia.

38= massulae 0=absent 1=orchidoid 2=epidendroid 3=arethusoid

Sectile pollinia have been shown to vary with respect to layering and regularity of massulae (Vermeulen, 1965; Freudenstein & Rasmussen, 1996). Orchidoid sectile pollinia have a single layer of uniform massulae, while epidendroid pollinia have variable numbers of layers of irregular massulae. Pollinia of Arethusa and Calopogon are hollow at maturity (cf. Pace, 1909; Freudenstein & Rasmussen, 1996), while most other orchids have solid pollinia.

39= pollinium texture 0=loose 1=hard

Pollinium texture is largely a continuous character (Dressler, 1986a), but it is possible to distinguish those pollinia that are truly coherent from those that are soft enough to be easily crushed when touched. The structural basis for this difference has been elucidated by ultrastructural studies of pollinia (Chardard, 1958, 1969; Cocucci & Jensen, 1969;Schill & Pfeiffer, 1977; Wolter & Schill, 1986; Yeung, 1987; Hu & Yang, 1989; Zavada, 1990; Pandolfi et al., 1993). The most important difference is whether or not exine is deposited on internal pollen grains; if not, a more cohesive, calymmate pollinium results, while those that do have exine on all grains are termed acalymmate and are much more friable (van Campo & Guinet, 1961).

40= poll. no.: 2 0=absent 1=present

41= poll. no.: 8 0=absent 1=longitudinal 2=transverse

The primary pollinium numbers in orchids are 2, 4, and 8. Other numbers are sometimes reported (e.g., in Laeliinae), depending upon whether additional small masses of pollen that sometimes are found along the caudicles are interpreted as pollinia. Four is the predominant number, and is found in the putatively basal orchid groups, as well as in outgroups (where there are four anther locules). Freudenstein & Rasmussen (1996) found that there are at least two ways to produce eight pollinia -- by longitudinal or transverse division of embryonic pollen masses.

42= pollinium orientation 0=juxtaposed 1=superposed

Variation in pollinium orientation was first recognized by Dressler & Dodson (1960). They described the superposed state, where, when four, the pairs of pollinia are stacked one on another, as opposed to the juxtaposed (Freudenstein & Rasmussen, 1996) condition, in which the pollinia are arranged side by side. We have detected two distinct ways in which the superposed state can occur -- either by inward or outward "rotation" of anther thecae (Freudenstein and Rasmussen, unpubl.), but as we have not yet been able to complete developmental study on a sufficient number of taxa, we have here coded all superposed pollinia as the same state.

43= ovary locule number 0=one 1=three

Most orchids have a single ovary locule, while a very few putatively basal groups have three locules, as does the outgroup. Transverse sections of each type are shown in Atwood (1984) and, in diagrammatic form, in Garay (1960).

44= stigma 0=convex 1=concave

Variation in stigma morphology was discussed by Dressler (1981, 1993), Rasmussen (1982) and Dannenbaum et al. (1989). In many taxa portions of one or more stigma lobes are protruded, form a raised triangular or circular mass, or are of other shape (cf. Rasmussen, 1982, fig. 73). These morphologies are grouped here under the term "convex". A concave stigma is a sunken, usually circular depression, which, as Dressler (1993) has suggested, appears to be specialized to receive hard pollinia (cf. Rasmussen, 1982, fig. 73:2.1.2); this type is usually encountered in Epidendroideae.

45= stigma receptive cells 0=various other 1=finger 2=prosenchymatic

This character derives from the work of Dannenbaum et al. (1989). It describes the shape of the receptive cells of the stigma that are usually hidden under stigmatic secretions in living plants.

46= viscidium 0=none 1=diffuse 2=detachable

Most orchids have some sort of adhesive associated with insect transfer of pollen masses. This may be either in the form of a glue that is transferred to the insect before it contacts pollen (diffuse), or a more elaborate cellular structure composed of rostellar tissue that is attached either directly, or via a stalk, to the pollinia (detachable). Some controversy over terminology involving the viscidium exists, with Dressler (1986a) and Dressler & Salazar (1991) arguing for restricting use of the term to a detachable structure, while Rasmussen (1982) used the term more broadly to include also any secreted adhesive. Here it is used in the broad sense simply for convenience, with the two senses of the term being the states. The character is coded as ordered because all detachable viscidia also have adhesive secretion.

47= endocarpic trichomes 0=absent 1=present

Endocarpic trichomes, or Schleuderhaare, were described by Beer (1857), and have been little discussed since. Hallé (1986) illustrated them in transverse sections of ovaries. An SEM of a hair from Pteroceras appears in Pedersen (1993). Their function is suspected to be in seed dispersal. Variation in shape (hairlike or flattened) occurs among genera. According to Malguth (1901), their presence is correlated with epiphytism, although not all epiphytes have them and some terrestrial orchids do. The only genus in which we found them outside of Epidendroideae is Prasophyllum.

48= seed lat. compressed walls 0=absent 1=present

49= seed testa cell shape 0=all iso 1=end iso, med elong 2=all elong

50= seed striations 0=absent 1=trans/ret 2=longitudinal

51= seed intercellular spaces 0=absent 1=present

52= seed wax caps 0=absent 1=present

53= seed covered cell border 0=absent 1=present

External morphology of seeds has provided a promising new set of data for orchids (Barthlott, 1976; Dressler, 1986b, 1990a, 1993; Molvray & Kores, 1993) These characters derive from the work of Ziegler (1981). Much of the scoring was done from the plates in Ziegler (1981), Tohda (1983, 1985, 1986), Chase & Pippen (1988, 1990), and Kurzweil (1993). Rather than coding seed morphology as types, such as those recognized by Ziegler (1981) and Dressler (1993), we coded component features to the extent possible. Laterally compressed testa walls refers to the extremely narrow cell lumen seen in some taxa (e.g., Vandeae), resulting from the close positioning of lateral anticlinal testa walls. Some seeds have distinct variation in testa cell size depending on location -- with either all cells isodiametric, the cells at both ends isodiametric and those in the center elongate, or all cells elongate (unordered). When present, striations on the sunken testa cell lumina may be either transverse/reticulate or straight-longitudinal (unordered). Distinct spaces occur among the cells in some taxa. "Wax caps" are present at the ends of testal cell protrusions in some members of Cymbidieae (Ziegler, 1981; Chase & Pippen, 1990). In some seeds the abuttment of adjacent testal cell walls is clearly evident, while in others, the line of demarcation between them is covered by tissue from one or the other cell (a covered cell border).


List of genera of Orchidaceae from the The Kew World Checklist of Monocotyledons (slightly modified), number of species in parentheses, dna available or obtainable, and number of species to sample.

This is also available as a searchable hyperbolic tree, but be sure to check the operating systems and browsers that are supported on the Phylogenetics page. TAKE A LOOK AT ALL OF THE GENERA OF ORCHIDACEAE ARRANGED BY SUBTRIBE GENERA OF ORCHIDACEAE

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ORCHIDACEAE (24,888 species)Tribes or subtribes (# of species)Genera (# of species)# to sample
Subfamilies (# of species) * = no DNA currenlty in hand, but if a number is in next column, we can get DNA of this genus 
APOSTASIOIDEAE (16) Apostasia (7)2
  Neuwiedia (9)1
VANILLOIDEAE (249)Pogoniieae (77)Cleistes (63)3
  Duckeella (3)1
  Isotria (2)1
  Pogonia (7)2
  Pogoniopsis* (2)0
 Vanillineae (172)Clematepistephium (1)1
  Cyrtosia (5)1
  Dictyophyllaria* (1)0
  Epistephium (23)2
  Eriaxis (1) 1
  Erythrorchis (3) 1
  Galeola (7)1
  Lecanorchis (14) 1
  Pseudovanilla (8)1
  Vanilla (109)11
CYPRIPEDIOIDEAE (154) Cypripedium (50) 7
  Mexipedium (1) 1
  Paphiopedilum (77) 25
  Selenipdeium (5)1
  Phragmipedium (21) 8
 Cranichideae (1643) 0
 Goodyerinae (651) Aenhenrya* (1) 0
  Anoectochilus (45)2
  Aspidogyne (19) 2
  Chamaegastrodia* (5) 0
  Cheirostylis* (47) 0
  Cystorchis* (21) 0
  Danhatchia (1) 0
  Dossinia (1) 2
  Erythrodes (88) 8
  Eurycentrum* (7) 0
  Evrardianthe* (1) 0
  Gonatostylis* (2) 1
  Goodyera (96) 10
  Gymnochilus* (3) 0
  Halleorchis (1) 0
  Herpysma* (1) 0
  Hetaeria (32) 2
  Hylophila* (10) 0
  Kreodanthus* (6) 2
  Kuhlhasseltia* (12) 2
  Lepidogyne* (1) 0
  Ligeophila (9) 1
  Ludisia (1) 1
  Macodes* (9) 2
  Microchilus (2) 0
  Moerenhoutia* (13) 0
  Myrmorchis (16) 0
  Odontochilus (3) 0
  Orchipedum* (2) 0
  Pachyplectron (3) 1
  Papuaea* (1) 0
  Platylepis (8) 1
  Platythelys (10) 2
  Pristiglottis* (22) 1
  Rhamphorhynchus* (1) 0
  Rhomboda (19) 0
  Stephanothelys* (2)0
  Tubilabium* (13) 0
  Vrydagznea* (41) 2
  Zeuxine (76)2
 Spiranthinae (470)Aracamunia* (1) 0
  Aulosepalum (6) 2
  Beloglottis (7) 1
  Brachystele* (20)1
  Buchtienia* (3) 0
  Coccineorchis (4) 1
  Cotylolabium (1) 0
  Cybebus* (1) 0
  Cyclopogon (82) 8
  Degranvillea* (1)0
  Deiregyne (19) 2
  Dichromanthus (1) 1
  Discyphus* (1) 0
  Eltroplectris (13) 0
  Eurystyles (18) 2
  Funkiella (4) 0
  Hapalorchis* (8) 0
  Helonoma* (2) 0
  Kionophyton (4) 0
  Lankesterella (11) 2
  Lyroglossa* (2) 0
  Mesadenella (7) 0
  Mesadenus (6) 1
  Microthelys (6) 0
  Odontorrhynchus (6)0
  Pelexia (77) 3
  Physogyne* (3) 0
  Pseudogoodyera* (2)0
  Pteroglossa* (9) 0
  Sacoila (5) 1
  Sarcoglottis (41) 4
  Sauroglossum* (12) 1
  Schiedeella (21) 2
  Skeptrostachys* (12) 0
  Spiranthes (24) 4
  Stalkya (1) 0
  Stenorrhynchos (7) 2
  Stigmatosema* (12) 0
  Svenkoeltzia (2) 0
  Thelyschista (1) 0
  Veyretia (9) 0
  Wallnoeferia* (1)0
 Manniellinae (1)Manniella (1)1
 Cranichidinae (268)Aa (27) 2
  Altensteinia (7) 2
  Baskervilla* (12) 2
  Coilochilus (1) 1
  Cranichis (54) 5
  Exalaria (1) 0
  Fuertesiella* (1) 0
  Gomphichis* (25) 3
  Myrosmodes* (10) 2
  Nothostele* (1) 0
  Ponthieva (57) 5
  Porphyrostachys* (2) 1
  Prescottia (31) 3
  Pseudocentrum* (7) 0
  Pseudocranichis* (1) 1
  Pterichis* (20) 2
  Solenocentrum* (4) 0
  Stenoptera (7)1
 Pterostylidinae (160)Pterostylis (160)16
 Chloraeinae (84)Bipinnula (11) 1
  Chloraea (51) 5
  Gavilea (14) 2
  Geoblasta (2) 0
  Megastylis (in part) (6)2
 Galeottiellinae (3)Galeottiella (3)1
 Diurideae (874) 0
 Acianthinae (170)Acianthus (26) 2
  Corybas (125) 12
  Cyrtostylis (6) 1
  Stigmatodactylus (11) 1
  Townsonia (2)0
 Caladeniinae (281)Adenochilus (2) 1
  Aporostylis (1) 1
  Caladenia (250) 25
  Cyanicula (10) 1
  Elythranthera (2) 1
  Eriochilus (12) 1
  Glossodia (2) 1
  Leptoceras (1) 1
  Praecoxanthus (1)0
 Cryptostylidinae (26)Coilochilus (1) 1
  Cryptostylis (25)3
 Thelymitrinae (166)Arthrochilus (10)0
   Burnettia (1)0
  Caleana (1) 1
  Calochilus (23) 5
  Chiloglottis (27) 5
  Drakaea (9) 3
  Leporella (1)1
  Lyperanthus (2)1
  Megastylis (in part) (1) 1
  Paracaleana (9)0
  Pyrorchis (2) 0
  Rimacola (1)1
  Spiculaea (1) 1
  Thelymitra (78)8
 Diuridinae (61)Diuris (58 6
  Epiblema (1)0
  Orthoceras (2)1
 Prasophyllinae (168)Genoplesium (45) 5
  Microtis (18) 2
  Prasophyllum (105)12
 Rhizanthellinae (2)Rhizanthella (2)1
 Codonorchideae (2)Codonorchis (2)1
 Orchideae (2185) 0
  Ponerorchis (17)2
 Brownleeinae (83)Brownleea (7) 2
  Disperis (76)2
 Disinae (220)Ceratandra (6)0
  Corycium (15)2
  Disa (169) 4
  Evotella* (1) 0
  Huttonaea (5) 1
  Pterygodium (18) 2
  Schizodium (6)0
 Orchidinae (1865)Aceratorchis* (2) 0
  Amerorchis (1) 1
  Amitostigma* (28) 2
  Anacamptis (13) 3
  Androcorys* (6) 1
  Aorchis* (2) 0
  Arnottia* (4) 0
  Barlia (2) 2
  Bartholina* (2) 1
  Benthamia* (31) 2
  Bonatea (17) 2
  Brachycorythis (36) 3
  Centrostigma* (3) 0
  Chamorchis (1) 1
  Chusua (20) 0
  Comperia (1) 1
  Cynorkis (158) 16
  Dactylorhiza (48) 5
  Diphylax* (4) 1
  Diplomeris* (5) 0
  Galearis (1) 1
  Gennaria (1) 1
  Gymnadenia (24) 2
  Habenaria (848) 85
  Hemipilia (18) 2
  Herminium (28) 2
  Himantoglossum (7) 2
  Holothrix (44) 3
  Megalorchis* (1) 0
  Neobolusia* (4) 0
  Neotinea (4) 1
  Neottianthe (8)1
  Ophrys (60)6
  Orchis (27)5
  Pachites* (2)0
  Pecteilis (4)1
  Peristylus (102)11
  Physoceras* (11)0
  Platanthera (135)14
  Platycoryne* (17)0
  Porolabium* (1)0
  Pseudorchis (1)1
  Roeperocharis* (5)0
  Satyrium (82)8
  Schizochilus* (11)2
  Serapias (14)2
  Smithorchis* (1)0
  Stenoglottis (4)2
  Steveniella (1)1
  Symphyosepalum* (1)0
  Thulinia* (1)0
  Traunsteinera (2)2
  Tsaiorchis* (1)0
  Tylostigma* (8)0
  Claderia* (2)1
 Neottieae (191)Aphyllorchis* (21)2
  Cephalanthera (20)2
  Epipactis (64)2
  Limodorum (3)2
  Neottia (63)6
  Palmorchis (20)2
 Sobralieae (237)Elleanthus (106)12
  Epilyna* (3)2
  Sertifera* (8)3
  Sobralia (120)15
 Tropidieae (35)Corymborkis (6)2
  Tropidia (29)2
 Triphoreae (28)Diceratostele (1)1
  Monophyllorchis (2)2
  Psilochilus* (7)1
  Triphora (18)2
 Nervilieae (73)Nervilia (65)2
  Epipogium (4)1
  Silvorchis* (1)0
  Stereosandra* (1)0
  Xerorchis (2)]1
 Gastrodieae (70)Auxopus (3)0
  Didymoplexiella (6)0
  Didymoplexis (17)2
  Gastrodia (41)4
  Neoclemensia* (1)0
  Uleiorchis (2)0
 Calypsoeae (70)Aplectrum (1)1
  Calypso (1)1
  Changnienia (1)1
  Corallorhiza (11)2
  Cremastra (3)1
  Dactylostalix (1)1
  Didiciea (2)0
  Ephippianthus (2)1
  Govenia (19)2
  Oreorchis (18)2
  Tipularia (5)2
  Yoania* (3)1
  [Wullschlaegelia (3)]1
  Chysis (8)2
  Coelia (5)2
 Ponerinae (22)Helleriella (2)1
  Isochilus (12)2
  Ponera (8)2
 Bletiinae (48)Basiphyllaea (7)2
  Bletia (34)4
  Hexalectris (7)2
 Pleurothallidinae (3999)Acianthera (131)15
  Anathallis (89)10
  Andinia (24)4
  Anthereon (6)2
  Barbosella (18)2
  Brachionidium (65)7
  Chamelophyton (1)0
  Dilomilis (5)1
  Diodonopsis (5)1
  Dracula (111)12
  Dresslerella (9)4
  Dryadella (42)4
  Echinosepala (8)2
  Frondaria (1)1
  Lepanthes (931)90
  Lepanthopsis (38)5
  Masdevallia (507)50
  Myoxanthus (44)10
  Octomeria (143)10
  Phloeophila (14)0
  Platystele (91)10
  Pleurothallis (813)80
  Pleurothallopsis (30)5
  Porroglossum (34)5
  Restrepia (48)5
  Restrepiella (1)1
  Scaphosepalum (41)5
  Specklinia (90)10
  Stelis (490)50
  Teaguia (10)10
  Tomzanonia (1)0
  Trichosalpinx (122)15
  Trisetella (22)5
  Zootrophion (12)5
 Laeliinae (1788)Acrorchis (1)1
  Alamania (1)1
  Arpophyllum (4)1
  Artorima (1)1
  Barkeria (15)3
  Brassavola (20)5
  Broughtonia (6)2
  Cattleya (54)5
  Caularthron (4)2
  Dimerandra (6)2
  Dinema (2)1
  Domingoa (2)1
  Encyclia (154)15
  Epidendrum (1125)112
  Euchile (2)2
  Hagsatera (2)0
  Hexisea (5)2
  Homalopetalum (4)2
  Isabelia (1)1
  Jacquiniella (6)2
  Laelia (11)2
  Lanium (9)2
  Leptotes (6)2
  Loefgrenianthus (1)0
  Meiracyllium (2)1
  Myrmecophila (10)2
  Nageliella (2)1
  Nanodes (2)1
  Neocogniauxia (2)1
  Nidema (2)1
  Oerstedella (29)5
  Oestlundia (11)2
  Orleanesia (9)0
  Pinelia (4)0
  Platyglottis (1)1
  Prosthechea (93)10
  Pseudolaelia (8)2
  Psychilus (15)2
  Pygmaeorchis* (2)0
  Quisqueya (4)1
  Renata (1)1
  Rhyncholaelia (2)2
  Scaphyglottis (63)6
  Schomburgkia (15)2
  Sophronitis (57)12
  Tetramicra (14)2
 Podochileae(1232) 0
 Eriinae (725)Ascidieria (1)1
  Ceratostylis (145)15
  Cryptochilus* (4)0
  Epiblastus (22)2
  Eria (404)4
  Mediocalcar (24)2
  Porpax (13)0
  Pseuderia (19)0
  Sarcostoma (5)2
  Stolzia (15)2
  Trichotosia (73)2
 Podochilinae (208)Appendicula (142)14
  Chitonochilus* (1)0
  Poaephyllum* (6)1
  Podochilus (59)2
 Thelasinae (299)Chitonanthera* (24)0
  Octarrhena* (41)0
  Phreatia (201)2
  Rhynchophreatia (9)1
  Ridleyella (1)0
  Thelasis (23)1
 Arethuseae (701) 0
 Arethusinae (9)Anthogonium (1)1
  Arethusa (1)1
  Arundina (1)1
  Calopogon (5)2
  Eleorchis (1)0
 Coelogyninae (692)Bletilla (5)2
  Bracisepalum (2)0
  Bulleyia (1)0
  Chelonistele (12)1
  Coelogyne (182)20
  Dendrochilum (264)28
  Dickasonia (1)0
  Dilochia (8)1
  Entomophobia (1)0
  Geesinkorchis (2)1
  Glomera (127)13
  Ischnogyne (1)0
  Nabaluia (3)1
  Neogyna (1)0
  Otochilus (5)0
  Panisea (8)1
  Pholidota (43)2
  Pleione (20)2
  Thunia (5)2
 Malaxideae (1158)Crossoglossa (21)2
  Hippeophyllum* (13)0
  Liparis (418)4
  Malaxis (395)4
  Oberonia* (308)30
  Orestias* (3)0
  Risleya* (1)0
 Cymbidieae (3814) 0
 Bromheadiinae (28)Bromheadia* (28)2
 Catasetinae (367)Catasetum (157)16
  Clowesia (7)3
  Cycnoches (33)5
  Cyrtopodium (44)4
  Dressleria (10)4
  Galeandra (34)2
  Grobya (4)2
  Mormodes (78)8
 Cymbidiinae (67)Cymbidium (51)12
  Grammatophyllum (11)2
  Graphorkis (4)2
  Porphyroglottis* (1)0
 Eulophiinae (316)Acriopsis (7)2
  Acrolophia* (7)0
  Ansellia (1)1
  Cymbidiella (3)2
  Cynaeorchis* (2)1
  Dipodium (24)2
  Eulophia (211)20
  Eulophiella* (5)2
  Geodorum* (13)2
  Grammangis (2)2
  Oeceoclades (38)2
  Thecopus* (2)0
  Thecostele (1)1
 Eriopsidinae (5)Eriopsis (5)2
 Oncidiinae (1589)Ada (16)5
  Amparoa (1)1
  Antillanorchis* (1)0
  Aspasia (7)7
  Brachtia (7)2
  Brassia (34)5
  Caluera (2)0
  Capanemia (14)2
  Caucaea (20)3
  Centroglossa (5)0
  Chytroglossa (3)0
  Cischweinfia (11)2
  Cochlioda (7)4
  Comparettia (7)3
  Cuitlauzina (1)1
  Cypholoron* (2)0
  Cyrtochiloides (3)3
  Cyrtochilum (119)20
  Diadenium* (2)2
  Dignathe (1)1
  Dunstervillea (1)0
  Eloyella (6)2
  Erycina (7)2
  Fernandezia (9)3
  Gomesa (17)2
  Goniochilus (1)1
  Hintonella (1)1
  Hofmeisterella (1)1
  Hybochilus (1)1
  Ionopsis (6)3
  Leochilus* (11)5
  Lockhartia (27)6
  Macradenia (11)2
  Macroclinium (38)4
  Mesospinidium (7)3
  Miltonia (10)4
  Miltoniopsis (5)3
  Neokoehleria* (10)2
  Notylia (58)6
  Odontoglossum (69)12
  Oncidium (336)50
  Ornithocephalus (44)8
  Osmoglossum (7)2
  Otoglossum (13)5
  Pachyphyllum (39)4
  Palumbina (1)1
  Papperitzia* (1)1
  Pfitzeria (1)1
  Phymatidium (8)2
  Platyrhiza (2)0
  Plectrophora* (10)2
  Polyotidium* (1)1
  Psychopsiella* (1)1
  Psychopsis (5)5
  Pterostemma* (2)1
  Quekettia* (7)0
  Raycadenco* (1)0
  Rodriguezia (48)5
  Rodrigueziella (6)2
  Rodrigueziopsis* (2)1
  Rossioglossum (4)2
  Rauhiella (3)0
  Rhynchostele (16)5
  Sanderella* (2)1
  Saundersia* (3)1
  Scelochiloides* (3)2
  Scelochilopsis (1)0
  Scelochilus* (48)8
  Seegeriella (1)1
  Sigmatostalix (56)8
  Solenidiopsis (4)2
  Solenidium* (2)2
  Stellilabium (34)5
  Stictophyllorchis* (2)0
  Stigmatorthos (1)0
  Suarezia* (1)1
  Sutrina* (1)1
  Symphyglossum (4)2
  Systeloglossum (5)2
  Telipogon (133)8
  Thysanoglossa (2)0
  Ticoglossum (2)2
  Tolumnia (36)10
  Trichocentrum (69)7
  Trichoceros (9)2
  Trichopilia (26)7
  Trizeuxis (1)1
  Warmingia (6)2
  Zelenkoa (1)1
  Zygostates (19)2
 Maxillariinae (749)Anguloa (11)2
  Anthosiphon (1)1
  Bifrenaria (26)16
  Chrysocycnis (4)2
  Cryptocentrum (19)6
  Cyrtidiorchis (4)1
  Horvatia* (1)0
  Ida (?)2
  Lycaste (50)5
  Maxillaria (552)60
  Mormolyca* (8)2
  Neomoorea (1)1
  Pityphyllum (5)5
  Rudolfiella (6)2
  Scuticaria (8)2
  Teuscheria* (7)2
  Trigonidium (14)4
  Xylobium (32)4
 Stanhopeinae (257)Acineta (15)5
  Archivea (1)0
  Braemia (1)1
  Cirrhaea (9)2
  Coryanthes (38)5
  Embreea (1)1
  Gongora (58)5
  Horichia (1)1
  Houlletia (9)2
  Kegeliella (3)2
  Lacaena (2)1
  Lueckelia (1)1
  Lueddemannia (1)1
  Paphinia (16)5
  Polycycnis (15)5
  Schlimmia (8)2
  Sievekingia (16)3
  Soterosanthus (1)1
  Stanhopea (55)5
  Trevoria (6)2
  Vasqueziella (1)1
 Coeliopsidinae (18)Coeliopsis (1)1
  Lycomormium (6)2
  Peristeria (11)2
 Zygopetalinae (418)Aganisia* (3)1
  Batemannia (5)2
  Benzingia* (2)1
  Bollea* (12)4
  Chaubardia* (5)5
  Chaubardiella* (8)5
  Cheiradenia* (1)0
  Chondrorhyncha (30)5
  Chondroscaphe (12)5
  Cochleanthes* (14)5
  Cryptarrhena (4)2
  Dichaea (110)25
  Dodsonia* (2)1
  Galeottia* (12)2
  Hirtzia (2)0
  Hoehneella* (1)0
  Huntleya* (13)4
  Kefersteinia* (61)8
  Koellensteinia (19)2
  Neogardneria* (1)0
  Otostylis* (4)1
  Pabstia* (6)1
  Paradisanthus* (4)1
  Pescatorea (16)4
  Promenaea* (19)2
  Stenia* (18)2
  Vargasiella* (2)0
  Warrea (4)2
  Warreella* (2)1
  Warreopsis* (4)1
  Zygopetalum (14)2
  Zygosepalum* (8)2
 Vandeae (2341) 0
 Polystachyinae (228)Hederorkis* (2)0
  Imerinaea* (1)1
  Neobenthamia (1)1
  Polystachya (224)25
 Aeridinae (1352)Abdominea* (1)1
  Acampe* (7)2
  Adenoncos* (16)2
  Aerides* (25)12
  Amesiella* (3)1
  Arachnis* (11)4
  Armodorum* (3)0
  Ascocentrum* (13)2
  Ascochilopsis* (2)1
  Ascochilus* (6)0
  Ascoglossum* (1)1
  Biermannia* (10)0
  Bogoria* (4)0
  Brachypeza* (7)0
  Calymmanthera* (5)0
  Ceratocentron* (1)0
  Ceratochilus* (2)1
  Chamaeanthus* (2)0
  Chiloschista* (20)2
  Christensonia (1)1
  Chroniochilus* (4)0
  Cleisocentron* (3)0
  Cleisomeria* (2)0
  Cleisostoma (87)9
  Cordiglottis* (7)0
  Cottonia* (1)0
  Cryptopylos* (1)0
  Dimorphorchis* (2)1
  Diplocentrum* (2)0
  Diploprora (2)1
  Dryadorchis (4)0
  Drymoanthus (4)0
  Dyakia (1)1
  Eparmatostigma (1)0
  Esmeralda* (2)1
  Gastrochilus* (55)6
  Grosourdya* (9)1
  Gunnarella* (9)0
  Haraella* (1)1
  Holcoglossum* (10)2
  Hygrochilus* (1)0
  Hymenorchis* (10)1
  Lesliea* (1)0
  Loxomorchis* (3)0
  Luisia* (39)2
  Macropodanthus* (6)0
  Malleola* (29)2
  Megalotus* (1)0
  Micropera* (18)2
  Microsaccus* (13)0
  Microtatorchis* (50)2
  Mobilabium* (1)0
  Neofinetia (2)1
  Nothodoritis* (1)0
  Omoea* (2)0
  Ornithochilus* (3)0
  Papilionanthe* (10)2
  Papillilabium* (1)0
  Paraphalaenopsis* (4)2
  Parapteroceras* (5)0
  Pelatantheria* (7)2
  Pennilabium* (10)0
  Peristeranthus* (1)0
  Phalaenopsis (62)12
  Phragmorchis* (1)0
  Plectorrhiza* (3)0
  Pomatocalpa* (33)4
  Porphyrodesme* (3)0
  Porrorhachis* (2)0
  Pteroceras* (24)2
  Renanthera* (17)2
  Renantherella* (2)0
  Rhinerrhiza* (1)0
  Rhynchogyna* (3)0
  Rhynchostylis* (3)2
  Robiquetia* (38)2
  Saccolabiopsis* (12)0
  Saccolabium* (10)0
  Sarcanthopsis* (1)0
  Sarcochilus* (25)2
  Sarcoglyphis* (12)1
  Sarcophyton* (3)1
  Schistotylus* (1)0
  Schoenorchis (26)*2
  Sedirea* (2)0
  Seidenfadenia* (1)1
  Smithsonia* (3)0
  Smitinandia* (3)1
  Staurochilus* (14)2
  Stereochilus* (7)2
  Taeniophyllum* (185)10
  Thrixspermum* (144)2
  Trichoglottis* (64)6
  Tuberolabium* (12)2
  Uncifera* (6)0
  Vanda* (57)6
  Vandopsis* (5)2
  Ventricularia* (2)0
  Xenikophyton* (1)0
 Angraecinae (445)Aeranthes (47)5
  Ambrella* (1)0
  Angraecum (219)22
  Bonniera* (2)1
  Calyptrochilum* (2)1
  Campylocentrum (73)5
  Cryptopus* (4)2
  Dendrophylax (9)5
  Harrisella (3)1
  Jumellea (58)2
  Lemurella* (4)1
  Lemurorchis* (1)1
  Listrostachys* (2)1
  Neobathiea* (5)2
  Oeonia (6)2
  Oeoniella (2)1
  Ossiculum* (1)0
  Sobennikoffia (4)2
 Aerangidinae (315) Aerangis (49)5
  Ancistrorhynchus* (16)2
  Angraecopsis* (21)2
  Beclardia* (1)1
  Bolusiella* (6)1
  Cardiochilos* (1)0
  Chamaeangis* (10)2
  Chauliodon* (1)0
  Cribbia* (4)1
  Cyrtorchis* (15)2
  Diaphananthe (24)2
  Dinklageella* (3)0
  Distylodon* (1)0
  Eggelingia* (3)0
  Eurychone* (2)1
  Margelliantha* (5)1
  Microcoelia* (29)2
  Microterangis* (7)1
  Mystacidium* (9)2
  Nephrangis* (2)1
  Plectrelminthus* (1)1
  Podangis* (1)1
  Rangaeris* (7)1
  Rhaesteria* (1)0
  Rhipidoglossum* (37)0
  Solenangis* (6)1
  Sphyrarhynchus* (1)1
  Summerhayesia* (2)0
  Taeniorrhiza* (1)0
  Triceratorhynchus* (1)0
  Tridactyle* (43)2
  Ypsilopus* (5)1
 Unplaced subtribes within Epidendroideae (3963) 0
 Agrostophyllinae (196)Adrorhizon* (1)1
  Aglossorhyncha* (13)1
  Agrostophyllum (91)9
  Earina (6)1
  Glossorhyncha* (80)0
  Ischocentrum* (2)0
  Sepalosiphon* (1)0
  Sirhookera* (2)1
 Dendrobiinae (3332)Bulbophyllum (1789)180
  Cadetia (60)2
  Chaseella* (1)0
  Dactylorhynchus* (1)0
  Dendrobium (1184)120
  Diplocaulobium (99)2
  Drymoda* (3)0
  Epigeneium (38)4
  Flickingeria (69)7
  Genyorchis* (7)2
  Jejosephia* (1)0
  Monomeria* (3)1
  Monosepalum* (3)0
  Pedilochilus* (35)0
  Saccoglossum* (5)0
  Sunipia* (22)2
  Trias* (12)2
 Collabiinae (435)Acanthephippium (12)2
  Ancistrochilus (2)2
  Aulostylis* (11)0
  Calanthe (187)2
  Cephalantheropsis* (5)1
  Chrysoglossum* (4)1
  Collabium (14)2
  Diglyphosa* (3)1
  Eriodes* (1)0
  Gastrorchis* (9)2
  Hancockia* (1)1
  Ipsea* (3)0
  Mischobulbum (9)2
  Nephelaphyllum (12)2
  Pachystoma* (1)1
  Phaius (48)2
  Plocoglottis* (39)2
  Spathoglottis (45)2
  Tainia (29)2
 Incertae sedis Thaia (1)0
 total ±2000

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Page last updated: 1 May 2013