Anatomy of the Monocotyledons Volume X: Orchidaceae

279 Pages·1960·45.93 MB·English

by  Cutler

ANATOMY OF THE
MONOCOTYLEDONS
x
ORCHIDACEAE
Anatomy of the Monocotyledons series
(Volumes I–VII edited by C. R. Metcalfe)
Vol. I. Gramineae. By C. R. Metcalfe, 1960.
Vol. II. Palmae. By P. B. Tomlinson, 1961.
Vol. III. Commelinales-Zingiberales. By P. B. Tomlinson, 1969.
Vol. IV. Juncales. By D. F. Cutler, 1969.
Vol. V. Cyperaceae. By C. R. Metcalfe, 1971.
Vol. VI. Dioscoreales. By E. S. Ayensu, 1972.
Vol. VII. Helobieae (Alismatidae) (including the seagrasses). By P. B. Tomlinson, 1982. ISBN 978–0–19–854502–6.
Vol. VIII. (eds M. Gregory and D. F. Cutler) Iridaceae. By Paula Rudall, 1995. ISBN 978–0–19–854504–0.
Vol. Ix. (eds M. Gregory and D. F. Cutler) Acoraceae and Araceae. By Richard C. Keating, 2002.
ISBN 978–0–19–854535–4.
Vol. x (eds M. Gregory and D. F. Cutler) Orchidaceae. By William Louis Stern, 2014. ISBN 978–0–19–968907–1.
Series editors: D. F. Cutler, M. Gregory and P. Rudall.
Anatomy of the Dicotyledons series
(Volumes I–III do not have editors)
Vol. I. Systematic anatomy of leaf and stem, with a brief history of the subject.
By C. R. Metcalfe and L. Chalk, 1979. ISBN 978–0–19–854383–1.
Vol. II. Wood structure and conclusion of the general introduction.
By C. R. Metcalfe and L. Chalk, 1983. ISBN 978–0–19–854559–0 (v. 1)
Vol. III. Magnoliales, Illiciales, and Laurales (Sensu Armen Takhtajan).
By C. R. Metcalfe, 1987. ISBN 978–0–19–854593–4 (v. 3).
Vol. IV. (eds D. F. Cutler and M. Gregory) Saxifragales.
By Hazel P. Wilkinson, R. J. Gornall, K. I. A. Al-Shammery and Mary Gregory, 1998. ISBN 978–0–19–854792–1.
Vol. I. reprinted with corrections in paperback, 1988. ISBN 978–0–19–854253–7.
Vol. II. reprinted with corrections in paperback, 1989. ISBN 978–0–19–854594–1.
Series editors: D. F. Cutler, P. Gasson, M. Gregory and D. W. Stevenson.
ANATOMY OF THE
MONOCOTYLEDONS
EDITED BY
M. GREGORY and D. F. CUTLER
Honorary Research Fellows,
Jodrell Laboratory, Royal Botanic Gardens,
Kew, Richmond, UK
x. ORCHIDACEAE
WILLIAM LOUIS STERN
Emeritus Professor of Botany,
University of Florida,
Gainesville,
Florida 32611, USA

INTRODUCTION
with an
by ALEC M. PRIDGEON
Sainsbury Orchid Fellow,
Royal Botanic Gardens,
Kew, Richmond, UK
1
1
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AUTHOR’S PREFACE
Sometime in the early 1980s I was asked by the late Charles Russell Metcalfe to prepare a compendium
on orchid anatomy along the lines of the already published plant families in Anatomy of the Monocoty-
ledons. Metcalfe, whom I admired, was Keeper of the Jodrell Laboratory at the Royal Botanic Gardens,
Kew, and a prominent plant anatomist. He was aware that many innovations had taken place during
the 50 or more years that had elapsed since a comprehensive account of orchid anatomy had appeared.
Technology had advanced appreciably and newly evolved ways of understanding systematics had been
applied to unravelling and answering botanical questions. Thus, the intent of this work is to provide an
up-to-date analysis of the vegetative anatomy of the orchid family (Orchidaceae), to suggest any taxo-
nomic implications of the application of anatomy to questions of systematics, and to present the results
in a systematic format providing a tool for systematists, students, and others to whom orchid anatomy
and systematics are relevant.
Heretofore, the only comprehensive body of work on orchid anatomy had been summarized by Hans
Solereder and Fritz Jürgen Meyer in their seminal compilation of 1930 as part of the series Systematische
Anatomie der Monokotyledonen. Solereder was Professor of Botany at the University of Erlangen, Ger-
many, and Meyer held a similar position at the Technical Institute in Braunschweig, also in Germany.
Solereder began the work and devoted more than 10 years to it. Besides compilations and interpreta-
tions of existing literature and his original observations, noted in the text by an exclamation point, ‘(!)’,
Solereder illustrated salient features. Following Solereder’s death in 1920, Meyer assumed editorship,
revising, updating, and completing Solereder’s work. The original product was published in German,
translated into English by A. Herzberg, and edited by B. Golek as part of the Israel Program for Scien-
tific Translation. The English version was published in Jerusalem in 1969.
Descriptions in Solereder and Meyer’s anatomical treatise were organ-based, however, making appli-
cation of their work cumbersome for systematic studies. There were no indices for taxa nor coordinated
lists of literature references; the bibliography was condensed into paragraph form and the entries were
listed chronologically rather than alphabetically by author. I have expanded the earlier efforts of Solered-
er and Meyer and presented results in a systematic format: by incorporating taxonomic categories, listing
literature and authors’ names alphabetically and chronologically, providing an evaluation for each taxon
described, and adding results from botanical literature and research since 1930. At the end of the intro-
ductory portion for each taxon described, I have added selected names of the chief ornamental species
represented in the taxon and listed species of putative medicinal value extracted from the compilations
of Lawler (1984), Gutiérrez (2010), and Singh and Duggal (2009).
It may be of interest that, prior to my study, an earlier unpublished work on the anatomy of orchids
had been attempted by Edward S. Ayensu while he was associated with the Smithsonian Institution. His
efforts lapsed owing to his feeling that he should put his energies into something else and to the lack of
collaborators who, almost correctly, considered the anatomy to be an endless task. There are estimated to
be more than 26, 000 species in the orchid family! I must confess that during my writing, at times, I was
reminded of the Herculean endeavours of James. A. H. Murray, compiler and editor of the great Oxford
English Dictionary, who spent 50 years on it! My own research began in 1983 while I was Professor and
Chairman of the Department of Botany at the University of Florida, Gainesville, and continued in
Florida after my 2002 retirement, studying at The Kampong, Coral Gables, and at Florida International
University, Miami and North Miami.
I have chosen to follow the format developed by Metcalfe in the Monocotyledon series he initiated,
including his anatomy of Gramineae in 1960 and, subsequently, anatomy of Cyperaceae in 1971. The
monocot series has continued, with the latest publication in 2002 being Volume Ix on Acoraceae and
Araceae by Richard C. Keating.
To provide a framework for this publication I have adopted the classification utilized in Genera Orchi-
dacearum (Pridgeon et al. 1999–2014). It is a fortunate circumstance that my work proceeds almost
simultaneously with the last volumes of Genera Orchidacearum. Resources for my work are my original
vi Author’s Preface
published research papers as well as publications of other botanists for groups unrepresented in my
studies.
I consider it an honour and pleasure to have been chosen to undertake this wholly enjoyable task. My
goals are for this treatise to enrich the body of information available on orchid anatomy, to serve as a
lodestone for future studies on this, arguably, the largest of flowering plant families, to provide a source
of taxonomic detritus to be picked over by systematists, and to unfold the rich fabric of orchid structure
as it applies to the biology of orchid life. My hope is to reach into the distant future with information we
need to sustain that which we know and to promote that which is waiting to be known about the most
seductive plants in the world.
W.L.S.
ACKNOWLEDGEMENTS
I have prefaced three volumes with the names of my authorities. I have done so because it is, in my opinion,
a pleasant thing and one that shows an honorable modesty to own up to those who were the means of one’s
achievements. . . .
Pliny, Natural History, Preface
It is an appealing thing for me to acknowledge the leadership and friendship of Charles Russell Metcalfe,
late Keeper of Kew’s Jodrell Laboratory, where, under his sponsorship, the seminal works Anatomy of the
Dicotyledons and Anatomy of the Monocotyledons were conceived and produced. I will feel everlastingly
honoured that he chose me to prepare this treatment of the orchids.
Every author needs an editorial companion to stand behind his/her shoulder; someone to comment
objectively, suggest revisions, search for and correct inaccuracies, understand the text, provide detailed
bibliographic commentary, guide the flowing and processing of words, and shadow his/her mental mus-
ings. Mary Gregory, Royal Botanic Gardens, Kew, has been that companion to me for which I will be
evermore grateful. Loismay C. Abeles, my wife and advisor on English didactics, has been with me
through writing the Preface, Acknowledgements, and sundry other parts of the text, bringing her skills
at composition and linguistic judgement to my aid. For her wisdom in these things and for standing by
me with her support I am supremely appreciative and thankful.
Alec M. Pridgeon, Royal Botanic Gardens, Kew, turned my interest from wood anatomy to orchid
anatomy. I sincerely thank him for moving me in that direction and for remaining my orchid mentor
these many years. Diurideae in the current work is the result of his study. For discussions on the cytology
of cells in apostasiad root tubercles, I acknowledge the interest and experience of Peter E. Brandham,
Royal Botanic Gardens, Kew, for confirming the presence of multinucleate cells in tissue sections of these
organs and for the use of his photomicroscope to picture them. My colleague, Barbara S. Carlsward,
Eastern Illinois University, acted as my anatomical barometer while she was my student exploring ideas
of cellular and tissue relationships and applying cladistic methods in our joint publications. She gracious-
ly provided illustrations of Vandeae from her study of this tribe for use in this work; additionally she
permitted me to use her unpublished observations on Sobralieae for this study.
Kurt Neubig, University of Florida, shared freely of his research on Sobralieae that formed the basis
of the current treatment of this tribe. I appreciate his generosity.
Beate H. E. Ruetter validated for me Max Siebe’s omission of root tubercles in his description of the
apostasias. Heike Hettler translated into English Martin Moebius’s German language monograph on
orchid leaf structure, for both of which I am grateful. Ruetter and Hettler served in my laboratory at the
University of Florida.
Over many years of friendship, the late Fred Fuchs, Naranja, Florida, kindly provided me with speci-
mens of uncommon orchid species for my research. Other major sources of research materials were
the Royal Botanic Gardens, Kew, through assistance there of Sandra Bell and the late Tim Lawrence. I
thank Ronald O. Determann, head conservator at the Atlanta Botanical Garden, for most of the vanilla
specimens I examined. It is a pleasure to acknowledge Kenneth M. Cameron, New York Botanical Gar-
den, for specimens of rare members of Vanilleae from the South East Asian islands without whose help
I would not have been able to report on the anatomy of these plants.
Words alone are often insufficient to express visual concepts. Pictures give life to words and David F.
Cutler, Royal Botanic Gardens, Kew, collaborated with Oxford University Press as liaison to organize
our pictures effectively, to evaluate the quality of the images, to oversee the preparation of captions, and
to collaborate with Mary Gregory in assembling the manuscript. I thank him for his diligence.
I thank James M. Heaney, Florida Museum of Natural History, for permission to use unpublished
material from university research on the anatomy of subtribe Polystachyinae, Beatrice N. Khayota, East
African Herbarium, National Museums of Kenya, for her study of Ansellia and related taxa, Marlene
viii Acknowledgements
Rosinski, Neuenkirchen, Germany, for her research analysis of Coelogyninae and Eriinae, and the late
Richard K. Baker, for leaf anatomy of Laeliinae, unpublished works from which I borrowed freely. There
were those who provided me with difficult-to-find plant material whom I wish to note with gratitude.
They are Brachycorythis from the late Joyce Stewart, Triphora from Susan M. Stern-Fennell, Coconut
Grove, Florida, and Calypso bulbosa from Kenton L. Chambers, Oregon State University, Corvallis.
Susan M. Stern-Fennell smoothed the way through customs at the airport in Miami for some of my field-
collected specimens. Some preserved specimens came from collections of Edward S. Ayensu, formerly
of the Smithsonian Institution. I have fond memories of Paul E. Stern, Ancile Gloudon, and Cicely
Tobisch, who accompanied me in Jamaica while I was collecting orchids for research; for their help in
the field I am grateful.
Wayne M. Morris, North Georgia College, Dahlonega, prepared most of the microscope slides of
tribe Dendrobieae and reported in his doctoral dissertation the anatomy of that group. I find pleasure in
acknowledging the skill of Lorraine M. McDowell, University of Florida, for preparing, processing, and
photographing tissues for both scanning and transmission electron microscopy.
Initially, my research took place at the University of Florida, Gainesville, where I was a faculty mem-
ber and chairman of the Department of Botany. I appreciate their generosity for providing laboratory
and office space and for making available equipment and supplies for my research.
Were it not for the help and influence of David W. Lee, Florida International University, I would not
have had laboratory, storage, and office space at Florida International University to continue research
after my retirement. I am grateful to him and to the university administration for accommodating me
and fulfilling my needs. The Kampong in Coral Gables, Florida, similarly provided space for my research
activities and writing.
Permission to use the line drawings of orchid tissues was granted by the Smithsonian Institution,
where I was chairman of the Department of Botany in the Museum of Natural History. These draw-
ings were executed by staff artists under the supervision of Edward S. Ayensu. The work was supported
through the Smithsonian’s Walcott Research Fund to which I express my thanks for making funds avail-
able. I also thank the owners and editors of the American Journal of Botany, the Botanical Journal of the
Linnean Society, the International Journal of Plant Sciences, Lankesteriana, and Lindleyana for permis-
sion to reproduce photographs from those publications.
For her expertise and knowledge of the intricacies of the computer, I must thank Vanessa Sandrino,
whose help smoothed the passage of text from mind to print.
I would be remiss were I not to acknowledge with thanks the technical expertise and services of the
librarians at the Royal Botanic Gardens, Kew, whose searches turned up many critical publications on
which some of the above descriptions and discussions are based.
I owe my deepest appreciation to the late Oswald Tippo, my guardian spirit, whose wise counsel
guided me safely through the meshwork of botanical idiosyncrasies, human and otherwise, and who
initially sparked my interest in plant anatomy.
CONTENTS
MATERIAL AND METHODS xi
INTRODUCTION by Alec M. Pridgeon 1
DESCRIPTIONS OF VEGETATIVE ANATOMY OF ORCHIDACEAE 43
Subfamily Apostasioideae 43
Subfamily Cypripedioideae 49
Subfamily Orchidoideae 52
Tribe Diseae 52
Tribe Diurideae by Alec M. Pridgeon 56
Subtribe Acianthinae 57
Subtribe Caladeniinae 58
Subtribe Cryptostylidinae 60
Subtribe Diuridinae 60
Subtribe Drakaeinae 61
Subtribe Megastylidinae 62
Subtribe Prasophyllinae 63
Subtribe Rhizanthellinae 64
Subtribe Thelymitrinae 64
Tribe Orchideae 65
Tribe Chloraeeae 73
Tribe Codonorchideae 73
Tribe Cranichideae 74
Subfamily Vanilloideae 79
Tribe Pogonieae 79
Tribe Vanilleae 81
Subfamily Epidendroideae 95
Tribe Arethuseae 95
Subtribe Arethusinae 95
Subtribe Coelogyninae 98
Tribe Calypsoeae 105
Tribe Collabieae 112
Tribe Epidendreae 116
Subtribe Bletiinae 116
Subtribe Chysinae 117
Subtribe Coeliinae 117
Subtribe Laeliinae 118
Subtribe Pleurothallidinae 130
Subtribe Ponerinae 135
Tribe Gastrodieae 136
Tribe Malaxideae 139
Tribe Neottieae 143
Tribe Nervilieae 148
Tribe Podochileae 150
Tribe Sobralieae 157
Tribe Triphoreae 161
Tribe Tropidieae 164
Tribe xerorchideae 168