List of reviewers

Demography and population dynamics

Flowering and survival of Dactylorhiza lapponica and Gymnadenia
in the Sølendet Nature Reserve, Central Norway
Dag-Inge Øien and Asbjørn Moen

A founder population of Orchis simia in The Netherlands:
a 30-year struggle for survival
Jo H. Willems

Dynamics of Orchis ustulata populations in Estonia
Kadri Tali

Temporal auto-correlation structures in populations of Cypripedium
: a two-year rhythm in flowering
Veijo Kaitala and Tiiu Kull

Dormancy and survival in rare terrestrial orchids
Richard P. Shefferson

Flowering and fruiting

Flower and fruit production in small populations of Orchis purpurea
and implications for management
Hans Jacquemyn, Rein Brys and Martin Hermy

Climatic influences on flowering and fruiting of Cypripedium
var. pubescens
Marilyn H. S. Light and Michael MacConaill

Key factors affecting shoot growth and flowering performance of
Dactylorhiza fuchsii
Petra Janecková and Pavel Kindlmann

Mycorrhiza and seed germination

Interactions between decomposing wood, mycorrhizas, and terrestrial
orchid seeds and protocorms
Dennis F. Whigham, Jay O’Neill, Melissa McCormick, Cynthia Smith,
Hanne Rasmussen, Bruce Caldwell and Tim Daniell

Germination of Dactylorhiza fuchsii seeds using fungi from non-orchid
Reem Salman, Grace Prendergast and Peter Roberts

The effects of fungicide and water application on seed germination and
infection in Gymnadenia conopsea under field conditions
Helen S. Scott and Peter D. Carey

Intensity of mycorrhizal infection in some orchid populations in Japan
Irina V. Tatarenko

Management and conservation

The sudden increase in the abundance of Himantoglossum hircinum in
England in the past decade and what has caused it
Peter D. Carey, Lynne Farrell and Nick F. Stewart

Population dynamics of Orchis morio in the Czech Republic under human
Jana Jersáková, Pavel Kindlmann and Michal Strítesky

Light climate and plant performance of Ophrys insectifera; a four-year
field experiment in The Netherlands 1998-2001
Edu Dorland and Jo H. Willems

An eleven year experimental study of the pink lady’s slipper orchid
Cypripedium acaule
Richard B. Primack

Species index

Subject index


The Orchidaceae contains the largest number of species of any family in the plant
kingdom (ca. 20 000 species) and it receives a large amount of attention because of
its enormous economic and social value compared to other plant families. From an
ecological perspective orchids are interesting because of their complex life cycle,
involving a vast array of pollination mechanisms and ubiquitous yet poorly understood
interactions with mycorrhiza. Orchids are also important from a biodiversity
perspective. They contribute large amounts of biodiversity to many ecosystems and
many species are recognized to be threatened or endangered. Although, most
orchids occur in the tropics as epiphytes, terrestrial species are widespread from the
subtropics to the arctic and they occur on all landmasses.
Many journals and books are devoted to orchids and there are orchid societies in
most countries, an expression of the broad public interest in the family. Some of the
attention placed on orchids is the result of the dramatic decline of species during the
course of the 20th century, mostly due to human activities such as harvest and habitat
modification. Only in recent years has attention focussed on orchid conservation
and restoration. Successful conservation and restoration requires knowledge of the
biology and ecology of orchid species, especially an understanding of the habitat
requirements of species and knowledge about ecological interactions between
orchids and fungi that are required by all orchids at some stage of their life cycle.
Conservation and restoration clearly needs to be based on knowledge of all aspects
of orchid biology, including heterotrophic and autotrophic stages of the life cycle.
The present book is a contribution toward a more complete understanding of the
biology and ecology of terrestrial orchids in the north temperate zone. It results
from a conference of specialists in the field of orchid biology and ecology in Ceské
Budejovice (Czech Republic) held from May 6-13th, 2001. The meeting was a follow-
up to a similar gathering that was held in The Netherlands in 1990 (Wells and
Willems 1991). The emphasis of the workshop in The Netherlands was on the population
ecology of terrestrial orchids, especially long-term demographic studies.
Other topics considered in the 1991 volume were patterns and causes of variation
in flowering and the use of models to project populations trends. The conference in
Czech Republic contained most of the themes presented at the 1990 conference but
expanded into areas that had previously received little attention. In particular, the
2001 conference included papers on orchid mycorrhiza, especially the role of mycorrhiza
on the underground phase (i.e., protocorms and seedlings) of orchids. Other
topics at the Ceské Budejovice meeting were seed germination and the effects of
site management on the population fitness.
A major shift in emphasis between the two conferences was an increase in the
number of experimental studies. Experimental studies, however need to be considered
as extensions of basic studies of orchid population biology and community
ecology because they lead to a better understanding of important elements of orchid
life cycles. There are, however, clear limits on the role of experimental studies of
endangered or threatened orchids. Many countries have restrictions, e.g experiments
must not be a threat for the populations under study. This often means that
research of the important underground phase is difficult to realize and, therefore
rather rare. Experimental approaches to the study of terrestrial orchids, both under
laboratory and field conditions, can be seen, however, as indispensable if we are to
unravel the mechanisms behind the population dynamics of terrestrial orchids.
Management will be most successful when conservation efforts are based on solid
scientific research.
The contributions in this volume are divided into four sections based on the general
themes of the papers. This does not mean, however that other aspects of orchid
ecology and biology can not be found in the several chapters. For instance, the first
three chapters also reflect, implicit or explicit, concern about the fate of the orchid
populations in nature. Hopefully, studies like the ones reported in this book will
contribute to the preservation of terrestrial orchids, which are not only flagship
species for nature conservation organizations, but also are indicative of environmental
We would like to thank the participants at the 2001 workshop for travelling to
the Czech Republic to attend the workshop. Their enthusiasm and efforts to compile
their presentations for inclusion in this volume are most appreciated. Many
interesting ideas were shared in Ceské Budejovice and we look forward to future
workshops and sharing of information.
We dedicate this book to the pioneer of modern orchid population research, Carl
Olof Tamm, Uppsala (Sweden), who started his ongoing studies already in the early
1940s and who contributed so much to the preservation of this peculiar plant family.
Literature cited
Wells, T.C.E. and J.H. Willems (eds.). 1991. Population ecology of terrestrial
orchids. SPB Academic Publishing bv, The Hague.

Pavel Kindlmann
Jo H. Willems
Dennis F. Whigham