Original Contributions to Botany, founded in 1886Edited by P. Leins, Heidelberg, H.-P. Comes, Salzburg, and S. Porembski, Rostock

ISSN 0067-7892

Volume 162

Schweizerbart Science Publishers

Dierk Michaelis

The Sphagnum Species of the World

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sBIBLIOTHECA BOTANICA, VOLUME 162

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sBIBLIOTHECA BOTANICA

Original Contributions to BotanyEdited by H. W. Lack, Berlin, P. Leins, Heidelberg and S. Porembski, Rostock

Volume 162

Dierk Michaelis

The Sphagnum Species of the Worldwith 219 plates and 15 text-fi gures

Schweizerbart Science PublishersStuttgart 2019

eschweizerbart_XXX

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sThe Sphagnum Species of the World

by

Dierk Michaelis1st english edition

with 219 plates and 15 fi gures

Schweizerbart Science PublishersStuttgart 2019

eschweizerbart_XXX

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sDierk Michaelis: The Sphagnum species of the world

Author’s address:Dr. Dierk MichaelisInstitute of Botany and Landscape EcologySoldmannstraße 1517487 Greifswald, Germany

We would be pleased to receive your comments on the content ofthis book:editors@schweizerbart.de

Introduction translated from the German by Philipp P. Thapa

Special thanks go to DUENE e.V. (part of the Greifswald Mire Centre) for granting a printing cost subsidy.

Front cover: Sphagnum pulchrum; Sweden; photo: D. Michaelis 2014

German edition: 978-3-510-48031-9 (Michaelis, Die Sphagnum-Arten der Welt)

ISBN: 978-3-510-48033-3ISSN: 0067-7892

Information on this title: www.schweizerbart.com/9783510480333

© 2019 E.Schweizerbart’sche Verlagsbuchhandlung (Nägele und Obermiller), Stuttgart, Germany

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical photocopying, recording, or otherwise, without the prior written permission of E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart

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Contents 1

Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 1.1. External and internal morphology of peat mosses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 1.2. Reproductive biology of peat mosses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 1.3. Research history . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 1.4. Evolutionary history . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 1.5. Ecology of the peat mosses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 1.6. Notes on identification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

2. Characteristic and key to sections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 2.1. Key to sections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 2.2. Short characteristic of sections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

3. Identification keys and regional species lists . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 3.1. Identification keys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 3.1.1. Europe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 3.1.2. Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 3.1.3. Africa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 3.1.4. North and Central America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 3.1.5. South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 3.1.6. Australia, New Zealand and Pacific . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 3.2. Regional species lists . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 3.2.1. Europe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 3.2.2. North Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 3.2.3. East Asia incl. Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 3.2.4. South Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 3.2.5. South-eastern Asia incl. New Guinea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 3.2.6. West Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 3.2.7. North Africa (Continent excl. the Azores) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 3.2.8. Central Africa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 3.2.9. South Africa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 3.2.10. Madagascar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 3.2.11 The Azores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 3.2.12. North America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 3.2.13. Central America continent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 3.2.14. Central America Antilles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 3.2.15. Northwestern South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 3.2.16. Eastern South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 3.2.17. Temperate South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 3.2.18. Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 3.2.19. New Zealand . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 3.2.20. Pacific . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71

4. Species descriptions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 4.1. Sphagnum sect. Acocosphagnum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 4.2. Sphagnum sect. Inretorta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

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4.3. Sphagnum sect. Sphagnum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74 4.4. Sphagnum sect. Rigida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89 4.5. Sphagnum sect. Insulosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91 4.6. Sphagnum sect. Acutifolia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132 4.7. Sphagnum sect. Squarrosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156 4.8. Sphagnum sect. Polyclada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157 4.9. Sphagnum sect. Acrosphagnum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207 4.10. Sphagnum sect. Subsecunda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211 4.11. Sphagnum sect. Isocladus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244 4.12. Sphagnum sect. Hemitheca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 331 4.13. Sphagnum sect. Cuspidata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 332 4.14. Sphagnum sect. Mollusca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 352 4.15. Ambuchanania . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 353

5. Glossarium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 399

6. References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 401

7. List of journals and monograph series . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 417

8. Illustration credits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 421

9. Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 425

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Preface 3

The Sphagnum or peat mosses are not only a species-rich genus. They influence like no other mosses the hydrology and hydrochemistry of their habitats, shape wide landscapes and sequestrate large amounts of car-bon in mires. Clymo & Heywood (1982) estimate, that Sphagnum comprises together more biomass than any other recent plant genus. Sphagnum dominated mires cover globally around two million square kilome-tres (Laine et al. 2009), and all mires contain at least 20 % of the global soil carbon (Maltby & Proctor 1996). The exceptional features of Sphagnum (water storage capacity, high productivity under nutrient-poor conditions, hardly to decompose under water-saturated conditions) give reasons for their large economic im-portance. They play a central role in the commercial horticulture in form of white peat and as biomass, are sometimes used as fuel or as sealing material. Due to its antiseptic effect, Sphagnum was used for wound band-age until the middle of the 20th century. Several projects of Sphagnum cultivation for sustainable utilisation are in well-advanced experimental phases (Gaudig 2002, Grantzau & Gaudig 2005, Pouliot et al. 2015).

The present work is an identification book for the species of the genus Sphagnum. It should enable the user to determine all Sphagnum species worldwide known so far. Concentration was put on the identifi-cation of the mostly vegetative material. This is the supplemented and reviewed version of the Sphagnum flora of Michaelis (2011), the first summarising global revision since Warnstorf’s Sphagnologia Universalis from 1911. There are not only approximately 150 spe-cies newly described since the time of Warnstorf, but also a lot of taxa revised and rejected to synonyms. The

revisions made by bryologist like Andrews, Isoviita, Eddy, Flatberg, and Crum form together with the new description mainly made by Crum, Flatberg, and An-drus another basis of this study. Furthermore, my own observation of Sphagnum mosses of by now five con-tinents are involved. Since time and means of transport are limited, it was not possible to study all Sphagna at their natural sites. Therefore, the results of literature research, the analysis of herbarium material and own observations are formed to one unit.

Towards the first edition, several recently described or revised species are involved and included in the keys. Furthermore, the introducing chapter was extended and reworked with regard to the phylogeny, the history of research and the ecology of Sphagnum.

I want to express special gratitude to the staff mem-bers of the herbaria of Berlin (FU), Hamburg, Geneva, Göttingen, Greifswald, Jena, London, Paris and Trond-heim, furthermore to the bryologists M. S. Ignatov (Moskau), A. I. Maksimov (Petrosavodsk) and O. L. Kusnezov (Petrosavodsk) for the procurement of not easily accessible Russian literature. Last, but not least, special thanks belongs to the colleagues of our work-ing group mire ecology of the Institute of Botany and Landscape Ecology (University Greifswald), who sup-plied me with numerous exotic Sphagnum specimens.

Finally, there is to say that in spite of multiannu-al work some questions are still open. Some species groups still need in-depth taxonomical analyses. How-ever, this book should support the Sphagnum identifi-cation in the scientific practice. Therefore, suggestions and constructive critiques for further improvements are always welcome.

Preface

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Abstract 5

Abstract

Sphagnum peat mosses are of immense economic and ecological importance. They occur on almost all conti-nents, mainly in northern South America, North Amer-ica and east and north-east Asia as well as in Europe.

The genus Sphagnum is very isolated within the Bryophyte plant division. Similarities in sporophyte design suggest a distant relationship with lantern moss-es (Andreaeopsida). Only recently the genus Ambucha-nania was identified as more closely related, but this genus similarly shows strongly derived characters. The genus Sphagnum is monophyletic; various approaches exist for its division in up to 4 subgenera and up to 18 sections. This revision discriminates 14 sections.

Among mosses the genus Sphagnum displays a singular combination of leaf-dimorphism (stem and branch leaves), branch dimorphism (spreading and hanging branches), and cell dimorphism (living chlo-rocytes and empty hyalocytes). Hence, assigning moss-es to the genus Sphagnum generally is not a problem. However, identification to the species level may be difficult because of the wide morphological plasticity in relation to hydrological conditions found in some species. In combination with differences in species concepts, this morphological plasticity has led to the recognition of narrowly or broadly defined species – up to the almost total negation of the existence of species. The total number of Sphagnum species has been esti-mated between 150 and 450.

This book offers an identification guide to all Sphagnum species worldwide. Since Carl Warnstorf’s

‘Sphanologia Universalis’, published in 1911 as part of A. Engler’s ‘Die natürlichen Pflanzenfamilien’, it is the first complete overview of the genus Sphagnum. Warnstorf used a rather narrow species concept, de-scribing many morphotypes as separate species. Since 1911, numerous synonyms have been recognised, par-ticularly through the works of Andrews, Eddy, and Iso-viita. Their revisions as well as c. 160 newly described species and own research provide the basis for the cur-rent volume. Delimitation of difficult taxa furthermore makes use of the results of recent genetic research.

Chapter 1 gives an overview of Sphagnum anatomy and morphology, providing the necessary termino logy for description and identification. In addition, the re-productive biology and phylogeny are described to-gether with a short account of research history. Chapter 2 provides descriptions of and a key to the sections. A key to the species, arranged by continent, is given in Chapter 3, complemented by species lists for 20 bioge-ographic regions. The keys to the Sphagnum species of Africa, Europe, and North America were developed on the basis of existing work. The keys for Asia and South America are entirely new. Detailed descriptions of 292 Sphagnum species can be found in Chapter 4. In addi-tion to morphological-anatomical characteristics, habi-tat, geographic distribution, and synonyms are given. This chapter includes 219 plates showing morphologi-cal and anatomical details. A comprehensive list of re-ferences allows for access to recent and older literature on Sphagnum.

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External and internal morphology of peat mosses 7

1.1. External and internal morphology of peat mosses

The overall shape of peat mosses can be characterised by the length and density of branches, the form of the head, and the growth direction. The growth is frequent-ly erect but can also be ascending or creeping. Young branches, which are short and densely grouped, form a more or less pronounced head at the apex of the plant, the so-called capitulum. In taxa with numerous branch-es, e.g. S. wulfianum, the capitulum is nearly spherical, otherwise hemispherical to flat. In specimens with re-duced branch formation, e.g. aquatic forms, the capitu-lum is absent. The characteristics of the capitulum thus depend on both the species and the environment.

The branches usually occur in bundles, the so-called fascicles. They are typically differentiated into pending and spreading branches. Pending branches have more imbricate and frequently more slender leaves and ap-pear thinner. Spreading branches have sometimes im-bricate, more often erect to spreading leaves and ap-pear thicker. This differentiation can be weak, as in some taxa of S. sect. Subsecunda, or fully absent, as in S. sericeum. Branches are mostly unramified. In a few taxa, one or two secondary branches originate near the base of the primary branch. The internal structure of the branches is usually differentiated into, from the outside in, a cortex, a sclerenchyma cylinder (or wood cylinder), and a parenchyma. The branch cortex is mostly single-layered. Reinforcing spiral fibres occur only in the section Sphagnum. The cells of the cortex can be monomorphic, as in the sections Sphagnum and Rigida, or differentiated into large cells with an apical pore, the so-called retort cells, and smaller cells with-out pores. An additional diagnostic trait is leaf arrange-ment. In most species, leaves are arranged helically. In some cases, the branch leaves are quinquefarious, i.e. ranked one after the next in five straight rows (S. quinquefarium, S. pentastichum = S. recurvum). The tips of the leaves can be imbricate, spreading, or squar-rosely curved.

In general, the branch leaves are relatively uniform. Their shapes range from ovate to lanceolate to almost awl-shaped. Branch leaves are usually concave with an incurved upper leaf surface. They mostly have a more or less narrow margin of undifferentiated cells. If the outer lateral wall of the rotund margin cells has been dissolved due to resorption, this is called a resorption furrow. It occurs in the sections Sphagnum and Rigida as well as in some Acutifolia species. The single-lay-ered lamina is almost always clearly differentiated into hyaline cells and chlorophyll cells. Hyaline cells are large and empty (dead). Chlorophyll cells are small, green, and arranged in a net pattern. The hyaline cells of the branch leaves are usually reinforced with fibrils and not divided lengthwise by septa. The fibrils can be shaped like thin rings or spirals, but they can also be flat (like a perforated disc) or band-shaped (‘meniscus-like’).

Pores in the outer walls of the hyaline cells facilitate water intake and discharge. In submerged peat mosses, pores are unnecessary and thus often reduced or absent.

Pores can be circular or elliptical, surrounded by a reinforced wall (ringed pores), and located in various

1. Introduction

Fig. 1: Branch cortex in various sections (1 – S. sect. Sphag-num, 2 – S. sect. Rigida, 3 – S. sect. Squarrosa, 4 – S. sect. Subsecunda, 5 – S. sect. Cuspidata, 6 – S. sect. Acutifolia)

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24 Introduction

to Michaelis (2011), the demarcation of regions was updated following Sjögren (2001), including a new demarcation between the Azores with their high di-versity of peat mosses from the low-diversity North African mainland. In addition, the distribution details incorporate information from the respective species de-scriptions, supraregional compendiums such as Cardot (1897), Herzog (1926), and Warnstorf (1894a, 1911, 1913) as well as regional floras. The European floras used are Blockeel (2011), Bouman (2002), Frey et al. (1995), Hill (2004), Ignatov & Ignatova (2003), Iso-viita (1970), Jóhannsson (1989), Laasimer et a. (1954), Laine et al. (2009, 2018), Proctor (1955), Savicz (1936), Séneca & Söderström (2009), Serov (1964), Vellak et al. (2013) , and Želesnova (1994). The Asian floras used are Abay & Keçeli (2014), Abramov & Abramova (1983), Afonina (2004, 2009), Bardunov (1965, 1974), Bardunov & Čerdanceva (1982), Bartram (1939), Czernyadjeva et al. (2009), Dixon (1935), Eddy (1977, 1988), Fedosov (2012), Hansen (1961, 1965), Igna-

tov & Afonina (1992), Iwatsuki (1991, 2004), John-son (1959, 1980), Kürschner et al. (2015), Maksimov (2007), Melin (1924), Muldijarov (1990), Pisarenko (2015), Redfearn et al. (1996), Stepanova (1986), and Suzuki (1955, 1972). The African floras used are Ah-Peng & Bardat (2005), Ah-Peng et al. (2010), Bates & Gabriel (1997), Born & Frahm (1993), Crosby et al. (1983), Dias et al. (2009), Eddy (1985, 1993), Frahm (2005), Frahm et al. (2009), Garside (1949), Magill (1981), Muller et al. (2010, 2011), Taylor & Thomp-son (1955), and Theriot (1922). The North American floras used are Allen (1994), Anderson (1990), Bartram (1949), Crosby (1969), Crum (1980a, 1984a, 1986), Duarte Bello (1997), McQueen & Andrus (2007), and Steere (1978). The South American floras used are Brotherus (1895), Churchill (1994, 1998), Churchill et al. (2000), Costa et al. (2011), Delgadillo et al. (1995), Florschütz-de Waard et al. (2011), Griffin (1981), Mit-ten (1869), Müller (2009), Pursell (1973), Ule (1899), Yano (1981, 1995, 1996), Yano et al. (1985), and Ziem-

Fig. 14: Distribution of Sphagnum species. The demarcation of regions follows Margadant & Florschütz (1967), adapted with modifi cations from Sjögren (2001). For regional inventories see Chapter 3.2.

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meck & Harpel (2012). The floras used on Australia and Oceania are Bartlett (1984), Beever et al. (1992), Fife (1995, 1996), Karlin & Andrus (1995), Scott & Stone (1976), Séneca & Söderström (2011), Seppelt (2000), Streimann & Curnow (1989), and Whinam et al. (2003).

Distribution data should be treated as preliminary. Current knowledge on the ranges of many tropical spe-cies remains insufficient, and even obvious mistakes in European distribution maps are not infrequent. (For example, the maps on S. angustifolium and S. fallax = S. recurvum var. mucronatum in Daniels & Eddy 1985 show large gaps in Central Europe.) For species with small (partial) ranges, geographical details were added. Information that bears limited relevance to spe-cies identification, such as spore size and colour or the number of chromosomes, is not included.

Ecological information has deliberately been kept short and should not be overestimated. While there are comprehensive publications on the autecology of

Fig. 15: Morphological variability of the multiform species Sphagnum denticulatum. All material from Sweden and Finland: A from mire ditch, B from a fl oating mat, C from a streamlet, and D from a lake bottom.

holarctic species, current knowledge on the needs of many tropical species is very poor. Especially older descriptions up to 1911 often include information on the provenance but not the habitat. Additionally, there are uncertainties in the use of some terms. The often used but rather loose concept of bog (including sites that range from strictly rainwater-fed raised bogs to any mire in which Sphagnum occurs) is rendered here as oligotrophic to mesotrophic acid mire. The nomen-clature of ecological mire types follows Succow & Joosten (2001). It should also be noted that the ecologi-cal needs of a species may vary within its distribution range (Gignac 1993) and are often unknown for a given location. While the habitat may serve as a clue to the identity of a species, such information cannot replace anatomical analysis.

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fibrils throughout and undivided, on ventral side with ringed, elliptic pores in cell angles and several smaller pores and pseudopores along the commissures, on dor-sal side with large, ringed pores in groups of three in adjacent cell angles and some single pores in lateral cell angles; branch fascicle: with 2 similar, short (about 5 mm long) and upwards oriented branches, cortex with delicate spiral fibrils and one pore per cell; branch leaf: elongated ovate with cucullate apex, 2.5–3.0 mm long, leaf margin with resorption furrow; branch leaf pores: pores similar to stem leaf pores; chlorophyll cells: in-verted narrow triangular, ventral exposed, dorsal barely enclosed by the hyaline cells or narrowly exposed; hya-line cells: ventral slightly convex, dorsal convex, walls adjacent to chlorophyll cells smooth.Habitat: submerged in hollows of blanket bogsDistribution: Am.4 (Venezuela: Estado Bolívar, Cerro Guaiquinima)

19. S. multifibrosum Xui J. Li & M. Zang ex Xui J. Li in Acta Bot. Yunnan. 6: 77. 1984 (Pl. 17)Habitus: robust to large plants; colour: pale green with yellow tinge; stem: cortex two- to three-layered, with spiral fibrils, on the external side with pores, wood cylinder yellow-brown; stem leaf: long lingulate, 3.8–4.3 mm long; hyaline cells: undivided or sparsely di-vided, with well developed fibrils and numerous ringed pores on the dorsal side; branch fascicle: with 4–5, thereof 2 spreading; branch leaf: roundish to ovate with cucullate apex, about 2.0 mm long, leaf margin with re-sorption furrow; branch leaf pores: on ventral side with a few ringed pores along the commissures, on dorsal side with ringed, elliptic pores along the commissures; chlorophyll cells: narrow inverted isosceles triangular, ventral exposed, dorsal enclosed; hyaline cells: ventral slightly convex to almost plane, dorsal convex, walls adjacent to chlorophyll cells smooth.Habitat: acid oligotrophic to mesotrophic mires, on moist sites in Picea forests and Rhododendron shrubs, on irrigated rocksDistribution: As.2 (China: southern Tibet to Fujian, Heilongjiang)

20. S. negrense Mitt. in J. Linn. Soc., Bot. 12: 624. 1869 (Pl. 16)Habitus: medium-sized to robust plants; colour: green, grey green, yellow green; stem: cortex two- to three-layered, without spiral fibrils, on the external surface mostly with one apical pore; stem leaf: ovate to ovate-lingulate with rounded, somewhat fringed and concave apex, 1.7–2.0 mm long, leaf margin with resorption furrow; hyaline cells: often one to several times di-

vided, with fibrils down to the leaf base, assemblage of pores similar to branch leaves; branch fascicle: with 2–3, thereof 1–2 spreading, cortex with few and very delicate spiral fibrils, mostly with one apical pore; branch leaf: ovate with cucullate apex; 1.4–1.5 mm long, leaf margin with resorption furrow; branch leaf pores: on ventral side in the upper half of leaf with lar-ger, weakly ringed pseudopores partially in rows along the commissures and with scattered very small, round, strongly ringed pores in cell angles or in the middle of hyaline cells, on dorsal side with numerous ellip-tic, ringed pores in loose rows along the commissures and also with 1–4 very small, strongly ringed pores in the middle of the hyaline cells in the upper half of leaf; chlorophyll cells: inverted triangular to trapezoid, ventral broadly exposed, dorsal enclosed, more rarely narrowly exposed; hyaline cells: ventral slightly con-vex, dorsal convex, walls adjacent to chlorophyll cells smooth.Remarks: The descriptions delivered by Warnstorf (1891, 1911) and Crum (1990, 1993) give no informa-tion about the differentiation and colour of the wood cylinder.Habitat: on moist stones near cataracts and water falls, in moist shrub vegetationDistribution: Am.4 (Colombia, Venezuela), Am.5 (Guayana, Brazil: Amazonas)

21. S. palustre L., Sp. Pl. 1106. 1753 (Pl. 18)Syn.: S. cymbifolium (Ehrh.) Hedw. 1782, S. deflexum Gilib. 1792, S. obtusifolium Ehrh. ex Hoffm. 1795, S. latifolium (Weiss) Hedw. 1802, S. vulgare Michx. 1803, S. oblongum P.-Beauv. 1805, S. crassisetum Brid. 1806, S. cymbifolioides Breut. 1824, S. cubile Neck. ex Brid. 1827, S. subbicolor Hampe 1880, S. glaucum Klinggr. 1880, ? S. puiggari Müll. Hal. 1887, ? S. brachybolax Müll. Hal. ex Warnst. 1891, ? S. suberythrocalyx Müll. Hal. ex Warnst. 1891, ? S. vitji-anum Schimp. in Warnst. 1891, S. japonicum Warnst. 1895, ? S. kegelianum Müll. Hal. ex Warnst. 1897, S. subtursum Müll. Hal. ex Warnst. 1897, ? S. brachycladum Müll. Hal. ex Warnst. 1897, S. heterophyllum Warnst. 1899, S. trachya-cron Müll. Hal. in Warnst. 1903, ? S. paranae Warnst. 1905, ? S. santosense Warnst. 1906, S. sulphureum Warnst. 1907, S. klinggraeffii Röll 1907, ? S. derrumbense Warnst. 1911, S. biforme Warnst. 1911, ? S. chlorocephalum Hampe in Warnst. 1911Habitus: mostly robust to large plants; colour: green, grey green, bluish green, yellow green to light brown-ish, at times flesh-coloured; stem: cortex mostly three-layered, with spiral fibrils, on the external surface with 1–4 (rarely up to 9) pores, wood cylinder yellow-brown to brown; stem leaf: lingulate to spatulate, 1.2–2.5 mm long; hyaline cells: undivided or sparsely divided,

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simplementation of fibrils and pores variable; branch fascicle: with 3–5, thereof 2–3 spreading; cortex with spiral fibrils; branch leaf: roundish to elongated ovate with cucullate apex, 1.7–2.8 mm long, leaf margin with resorption furrow; branch leaf pores: on ventral side with several roundish pores mostly along the commis-sures, on dorsal side with elliptic, ringed pores along the commissures; chlorophyll cells: narrow inverted isosceles triangular to trapezoid, ventral exposed, dor-sal enclosed or narrowly exposed; hyaline cells: ventral slightly convex, dorsal convex, walls adjacent to chlo-rophyll cells smooth.Habitat: often acid mesotrophic mires, up to acid eu-trophic mires (Alnus forests)Distribution: Eur., As.1, As.2, As.3, As.4, Afr.5 (the Azores), Am.1, Am.4, Am.5, Oc

22. S. pseudocymbifolium Müll. Hal. in Linnaea, 38: 547. 1874 (Pl. 19)Syn.: S. cymbifolium Mitt. 1859, S. assamicum Müll. Hal. 1887, S. siamense Dix. 1932.Habitus: robust plants; colour: light brownish; stem: cortex two- to four-layered, with spiral fibrils, on the external surface with 2–7 pores per cell, wood cylin-der dark brown; stem leaf: lingulate to spatulate, at the apical margin somewhat fringed, 1.4–1.5 mm long; branch fascicle: with 3–4, thereof 2 spreading, cortex with spiral fibrils; branch leaf: roundish to ovate with cucullate apex, about 1.7 mm long, leaf margin with resorption furrow; branch leaf pores: on ventral side with several roundish, non-ringed pores mostly along the commissures, on dorsal side several mostly ringed pores along the commissures; chlorophyll cells: invert-ed equilateral triangular (to somewhat trapezoid) (most important difference to S. palustre), ventral exposed, dorsal scarcely enclosed; hyaline cells: ventral convex, dorsal strongly convex, walls adjacent to chlorophyll cells smooth.Habitat: in grassland near streamlets, in acid oligo-trophic to mesotrophic mires, on wet sites in forests and Rhododendron-shrublandsDistribution: As.2, As.3

23. S. reclinatum H.A. Crum in Contr. Univ. Michigan Herb. 20: 132. 1995 (Pl. 20)Habitus: small plants with lying to ascending growth habit; colour: pale green; stem: cortex two-layered, without spiral fibrils, on the external surface with scattered single pores, wood cylinder yellow-brown; stem leaf: similar to branch leaf, elongated ovate, the upper half constricted to an obtuse, cucullate apex, the upper part more or less squarrose, 2.5–3.0 mm long;

hyaline cells: with fibrils, undivided, on ventral side with few or lacking pores, on dorsal side with elliptic pores, mostly in groups of three in adjacent cell angles; branch fascicle: single on the upper side of stem, cor-tex without spiral fibrils, mostly without pores, rarely with single pores; branch leaf: elongated ovate, the up-per half constricted to an obtuse, cucullate apex, the upper part more or less squarrose, up to 2.0 mm long, leaf margin with resorption furrow; branch leaf pores: pores similar to stem leaf pores; chlorophyll cells: nar-row inverted trapezoid, ventral broadly exposed, dorsal narrowly exposed; hyaline cells: ventral slightly con-vex, dorsal convex, walls adjacent to chlorophyll cells smooth.Habitat: on sandy soil along white water rivers in pri-mary forestDistribution: Am.4 (Venezuela: Amazonas)

24. S. simplicicaulis H.A. Crum in Contr. Univ. Michi-gan Herb. 17: 76. 1990 (Pl. 20)Habitus: small plants, branchless, rarely dichotomous branched; colour: pale brown; stem: cortex two-lay-ered, with delicate spiral fibrils, on the external surface with one pore per cell, wood cylinder dark brown; stem leaf: similar to branch leaves, from a narrowed base widened to the middle of the leaf, above with rounded, cucullate apex, 2.5–2.8 mm long; hyaline cells: well fibrillose, on ventral side with several round to elliptic pores of variable size along commissures and in cell angles, on dorsal side with ringed, elliptic pseudopores, partially with real pores, mostly in groups of three in adjacent cell angles; branch fascicle: with only scat-tered single branches, weakly differentiated; branch leaf: weakly differentiated, ovate with cucullate apex and somewhat narrower leaf base; branch leaf pores: pores similar to stem leaf pores; chlorophyll cells: narrow inverted triangular, ventral exposed, dorsal scarcely enclosed by the hyaline cells or very narrowly exposed; hyaline cells: ventral slightly convex, dorsal convex, walls adjacent to chlorophyll cells smooth.Remarks: There are some similarities between the sparsely branched peat mosses S. reclinatum, S. sim-plicicaulis and S. subsecundoides. Sphagnum subse-cundoides could be differentiated from the other species by the smaller and more numerous pores. Sphagnum simplicicaulis shows a dark brown wood cylinder and at least delicate spiral fibrils inside the stem cortex whereas Sphagnum reclinatum has no spiral fibrils and a yellow-brown wood cylinder.Habitat: on sandy soil near rapidsDistribution: Am.4 (Venezuela: Estado Bolívar, Chi-manta Massif)

Sphagnum sect. Sphagnum 81

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25. S. subsecundoides H.A. Crum & W.R. Buck in Brittonia 44: 453. 1992 (Pl. 20)Habitus: very small to small plants; colour: yellow-ish; stem: cortex two-layered, rarely with delicate spi-ral fibrils, on the external surface often with one apical pore, wood cylinder yellowish, weakly differentiated; stem leaf: ovate-lanceolate with cucullate apex simi-lar to branch leaves, concave, 2.2–3.0 mm long, leaf margin upwards with resorption furrow; hyaline cells: often one time divided, with fibrils throughout to the leaf base, on ventral side with a few (1–4) pseudopores in cell angels and along the commissures, more com-mon near the apex, on dorsal side with numerous rather small, elliptic, ringed pores in loose rows along the commissures, near the apex occasionally with medium-sized, round pores near the middle of the hyaline cells; branch fascicle: single branches, 5–6 mm long, cortex occasionally with delicate spiral fibrils, often with one apical pore; branch leaf: broadly ovate with constrict-ed, cucullate apex, 2.2–3.0 mm long, leaf margin with resorption furrow; branch leaf pores: on ventral side with few to numerous (2–11) pseudopores in cell an-gles und along the commissures, on dorsal side with numerous rather small, elliptic, ringed pores in loose rows along the commissures, near the apex occasion-ally with medium-sized, round pores near the middle of the hyaline cells; chlorophyll cells: narrow inverted triangular, ventral broadly exposed, dorsal enclosed; hyaline cells: on both sides convex, walls adjacent to chlorophyll cells smooth.Habitat: on shores of rivers in primary forestDistribution: Am.5 (Brazil: Amazonas)

Species group 5: Species with smooth, elliptic branch leaf chlorophyll cells

26. S. amazonicum H.A. Crum & W.R. Buck in Brit-tonia 44: 449. 1992 (Pl. 21)Habitus: rather small plants; stem: cortex two- to four-layered, without spiral fibrils, on the external surface often with one pore per cell, wood cylinder dark red brown; stem leaf: lingulate, 1.2 mm long; hyaline cells: undivided or only sometimes one time divided, with complete or rudimentary fibrils (resorption), on ven-tral side membrane mostly resorbed or with 1–2 vari-able shaped gaps, on dorsal side in the upper third with 3–5 large, elliptic, ringed pores in adjacent cell angles, occasionally also along the commissures, directly at the apex with membrane gaps; branch fascicle: with 3, thereof 2 spreading, cortex without spiral fibrils,

often with one pore; branch leaf: ovate with cucullate apex, 2.0 mm long, leaf margin with resorption furrow; branch leaf pores: on ventral side with 2–4 small, ellip-tic pseudopores along the commissures, on dorsal side with large, ringed pores in groups of three in adjacent cell angles and some scattered pores along the commis-sures; chlorophyll cells: elliptic, centred and on both sides enclosed by the hyaline cells, walls adjacent to chlorophyll cells smooth.Remarks: There are some similarities with respect to leaf anatomy to S. sanguinale and S. magellanicum. However, Sphagnum sanguinale shows one to sev-eral times divided stem leaf hyaline cells, often with-out pores and fibrils, whereas Sphagnum magellani-cum usually has spiral fibrils in the cortex of stem and branches.Habitat: not mentionedDistribution: Am.5 (Brazil: Amazonas)

27. S. amoenoides H.A. Crum in Contr. Univ. Michi-gan Herb. 21: 147. 1997 (Pl. 21)Habitus: small plants; colour: pale green with brownish tinge; stem: cortex without spiral fibrils, on the external surface without pores, wood cylinder brown; stem leaf: ovate to ovate-lingulate with rounded, slightly concave to flat apex, 1.8 mm long, leaf margin with resorption furrow; hyaline cells: with fibrils almost down to the leaf base, on ventral side without pores, on dorsal side with small, strongly ringed pores in cell angles; branch fascicle: with 3, thereof 2 spreading, cortex without spiral fibrils, branch leaves moist five-ranked; branch leaf: ovate-lanceolate with cucullate apex, 1.5–1.8 mm long, leaf margin with resorption furrow; branch leaf pores: on ventral side without pores, on dorsal side with small, strongly ringed pores in cell angles; chlo-rophyll cells: barrel-shaped, on both sides exposed; hyaline cells: on both sides convex, walls adjacent to chlorophyll cells smooth.Remarks: An isophyllous peat moss, which is es-pecially characterised by its small, strongly ringed pores.Habitat: not mentionedDistribution: Am.5 (Brazil: São Paulo)

28. S. aureum C.B. McQueen in Bryologist 92: 405. 1989 (Pl. 22)Habitus: robust plants; colour: pale gold-coloured, yel-low-brown or light brown; stem: cortex three-layered, with spiral fibrils, on the external surface with 1–2 (–3) pores per cell, the apical pores often transverse elliptic; stem leaf: lingulate, 1.0–1.4 mm long; hyaline cells: un-divided, without fibrils and pores; branch fascicle: with

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s4, thereof 2 spreading, cortex with spiral fibrils and one pore per cell; branch leaf: ovate with cucullate apex, 1.0–1.5 mm long, leaf margin with resorption furrow; branch leaf pores: on ventral side without pores or with only a few pores, on dorsal side with pores in groups of three in adjacent cell angles or in groups of two in neighbouring hyaline cells; chlorophyll cells: elliptic, centred and on both sides enclosed by the hyaline cells; hyaline cells: on both sides slightly convex to almost plane, walls adjacent to chlorophyll cells smooth.Habitat: on wet seeps and the edges of eroded stream banks in páramo vegetationDistribution: Am.2 (Costa Rica)

29. S. buckianum H.A. Crum in Bryologist 95: 419. 1992 (Pl. 22)Habitus: medium-sized to rather small plants; colour: light green; stem: cortex two- to three-layered, without spiral fibrils, on the external surface without pores or with one pore per cell; stem leaf: roundish ovate to lin-gulate, 1.5–1.7 mm long; hyaline cells: in the upper half of leaf undivided and with fibrils, on dorsal side with 3–4 roundish, very small pores in cell angles; branch fascicle: with 2, both spreading; branch leaf: ovate with cucullate apex, 1.5 mm long, leaf margin with resorp-tion furrow; branch leaf pores: on ventral side with 2–3 pores or pseudopores in cell angles or along the commissures, on dorsal side with large, elliptic, ringed pores, often in adjacent cell angles; chlorophyll cells: elliptic, centred and on both sides enclosed by the hya-line cells; hyaline cells: on both sides slightly convex, walls adjacent to chlorophyll cells smooth.Habitat: on moist slopesDistribution: Am.5 (Brazil: São Paulo)

30. S. centrale C.E.O. Jensen ex Arnell & C.E.O. Jen-sen in Bih. Kongl. Svenska Vetensk.-Akad. Handl. 21: 34. 1896 (Pl. 23)Syn.: S. subbicolor auct., non Hampe 1880, S. palustre subsp. intermedium Russow 1887, S. papillosum var. inter-medium (Russow) Warnst. 1891, S. intermedium (Warnst.) Russow 1894.Habitus: robust to large plants; colour: pale to yellow-ish-green or brownish; stem: cortex four- to five-lay-ered, with spiral fibril, on the external surface with 1–5 (–10) pores, wood cylinder yellow to brown; stem leaf: lingulate, 1.2–2.2 mm long; hyaline cells: mostly undi-vided, mostly without fibrils; branch fascicle: with 4–6, thereof 2–3 spreading, cortex with spiral fibrils; branch leaf: ovate with cucullate apex, about 1.7 mm long, leaf margin with resorption furrow; branch leaf pores: on ventral side with few pores, on dorsal side with sev-

eral medium-sized to large, partially ringed pores and pseudopores; chlorophyll cells: elliptic, centred or ap-proximated to the ventral side, on both sides enclosed by the hyaline cells or narrowly exposed by a thickened outer wall; hyaline cells: ventral slightly convex, dorsal convex, walls adjacent to chlorophyll cells smooth.Habitat: acid mesotrophic miresDistribution: Eur., As.1, As.2, Am.1

31. S. cristatum Hampe in Linnaea 38: 661. 1874 (Pl. 24)Syn.: S. pachycladum Müll. Hal. in F. Müll. 1882, S. leiono-tum Müll. Hal. 1887, S. trachynotum Müll. Hal. 1887, S. whiteleggei Müll. Hal. 1887, S. wilcoxii Müll. Hal. 1887, S. australe Schimp. in Warnst. 1890, S. maximum Warnst. 1891, S. pseudomedium Warnst. 1891, S. cymbophyllum F. Müll. in Warnst. 1891, S. microcephalum Müll. Hal. in Warnst. 1900, S. dielsianum Warnst. 1904, S. otagoense Warnst. 1904, S. wardellense Warnst. 1907, S. cymbophylloides Warnst. 1907, S. decipiens Warnst. 1907, S. maori-compactum Müll. Hal. in Warnst. 1911.Habitus: rather small to robust plants; colour: pale to dark brownish green, sometimes pale green, rarely purple-brown; stem: cortex three- to four-layered, with spiral fibrils, on the external surface with (1–) 2–5 round or elliptic pores, wood cylinder brown; stem leaf: lingulate, (1.1–) 1.5–2.2 mm long; hyaline cells: mostly undivided and with fibrils, on ventral side with few or without pores, on dorsal side with several large, ringed pores or, especially in the upper third, with large membrane gaps; branch fascicle: with (3) 4–6, thereof 2–3 spreading, cortex with spiral fibrils, partially with one apical pore; branch leaf: broadly ovate with cucul-late apex, 2.0–2.5 mm long, leaf margin with resorption furrow; branch leaf pores: on ventral side pores lacking or with scattered large pores, on dorsal side with 2–8 large, ringed pores, often in groups of three in adjacent cell angles; chlorophyll cells: elliptic, on both sides enclosed by the hyaline cells or narrowly exposed; hyaline cells: on both sides slightly convex to almost plane, walls adjacent to chlorophyll cells smooth.Habitat: acid oligotrophic mires, wet heath (Whinam et al. 2003)Distribution: Austr.1 (Mainland Australia, Tasmania), Austr.2 (New Zealand, the Chatham Islands), Oc. (New Caledonia)

32. S. cuculliforme H.A. Crum in Contr. Univ. Michi-gan Herb. 16: 141. 1987 (Pl. 25)Habitus: small plants; colour: at the head pink or pink-brownish, below pale to brownish white; stem: cortex two- to three-layered, without spiral fibrils, on the external surface often with one large, roundish pore at the apical end, wood cylinder dark reddish brown;

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206 Species descriptions / plate 89

Plate 89: Sphagnum wulfi anum: a: habitus, b: branch fascicle, c: stem cross-section, d: stem cortex, e: branch cortex, f: branch cross-section, g: stem leaves, h: upper stem leaf cells dorsal, i: branch leaves, j: branch leaf cells ventral, k: branch leaf cells dorsal, l–m: branch leaf cross-section.

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sface without pores, wood cylinder dark red to black red; stem leaf: triangular-lingulate with rounded and narrowly truncated, dentate or somewhat fringed apex, 0.5–0.8 mm long; border: narrow, downwards not wid-ened; hyaline cells: in the upper half of leaf mostly one time, occasionally two times divided, without fibrils, on both sides partly or largely resorbed; branch fasci-cle: with 6–13, thereof 3–5 spreading, cortex with re-tort cells; branch leaf: narrowly ovate-lanceolate with a long, involute, narrowly truncated and dentate leaf tip, 1.0–1.4 mm long, leaf margin without resorption furrow; branch leaf pores: on ventral side few small, unringed pores in cell angles, near leaf margins often few larger pores, on dorsal side with several small, ringed pores in cell angles and along the commissures in the upper half of leaf; chlorophyll cells: rectangular to barrel-shaped, in the leaf tip elliptic, centred, on both side with more or less thickened cell walls exposed, only in the upper third also enclosed by hyaline cells; hyaline cells: on both sides slightly convex to almost plane, walls adjacent to chlorophyll cells faintly to dis-tinctly papillose.Habitat: moist coniferous and mixed coniferous for-estsDistribution: Eur. (especially Northern Europe, rare in Eastern Europe), As.1, Am.1

4.9. Sphagnum sect. Acrosphagnum Müll.Hal. 1887

119. S. capense Hornsch. in Linnaea 15: 113. 1841 (Pl. 90–92)Syn.: S. mollissimum Müll. Hal. 1887, S. panduraefolium Müll. Hal. 1887, S. austro-molle Müll. Hal. 1887, S. subro-tundifolium Müll. Hal. 1900, S. wenckei Röll 1907, S. pap-peani Breutel in Warnst. 1911, S. humbertii Cardot 1916, S. bourbonense H.A. Crum 1992Habitus: small, slender plants; colour: in shadow green, elsewhere with orange-brown tinge or complete plants orange-brown, sometimes with reddish purple tinge; stem: cortex two- to three-layered (up to four-layered), on the external surface often with one large pore or without pores, wood cylinder yellow or brown; stem leaf: ovate-lingulate with broadly rounded, trun-cated and dentate, more or less concave apex, 1.2–1.9 mm long; border: narrow, not widened downwards; hyaline cells: undivided or rarely with scattered septa, often in the upper half of leaf with fibrils, sometimes with fibrils down to the leaf base, in the fibrillose part implementation of pores similar to branch leaves, on

ventral side with numerous commissural pores or only scattered pores in cell angles, on dorsal side in the up-per half mostly with many rather large, ringed pores along the commissures and membrane gaps of cell width, additionally often one to several centred pores, in the lower half only single large gaps in the middle of cells or in distal and proximal cell angles; branch fascicle: with 2–4 (–5), thereof 2 (–3) spreading, not or hardly dimorphous, cortex with often two retort cells, one after another, branch leaves more or less distinctly five-ranked; branch leaf: ovate to ovate-lanceolate with narrowly truncated leaf tip with 3–5 denticles, 0.7–1.0 mm long; branch leaf pores: variable, on ventral side with scattered pores in cell angles or in discon-tinuous rows along the commissures, occasionally with many pores in dense rows, on dorsal side often with many medium-sized, ringed pores in dense rows along the commissures and several, partly ringed, centred pores, rarely pores restricted to apical part, occasionally pores partly displaced by pseudopores; chlorophyll cells: ellip-tic, ventral often enclosed, dorsal narrowly exposed by thickened cell wall, more rarely on both sides exposed resp. enclosed; hyaline cells: on both sides slightly con-vex, walls adjacent to chlorophyll cells smooth.Remarks: According to Crum (1992a), S. bourbon-ense differs from S. capense by similarity of branch and stem leaves in shape and porosity, large and weakly ringed dorsal branch leaf pores (while ventral pores are few or lacking), very narrow branch leaf chlorophyll cells, and undivided stem leaf hyaline cells. Whereas the leaf shapes, the undivided stem leaf hyaline cells and the branch leaf cross-sections actually hardly dif-fer (see species diagnosis of S. capense in Warnstorf 1911), “typical” S. capense shows at least often some-what smaller (4–6 μm) and ringed dorsal commissural pores. The specimen of De Slover 17.625 from Ré-union most similar to S. bourbonense (in Eddy 1985, Fig 27C) is evaluated differently by Crum (1992a) and assigned to S. bourbonense. The examined specimens of S. bourbonense (paratype and isotype, JE 04006740 and JE 04006808) show on dorsal side often ringed (!) branch leaf pores of 7–11 μm in diameter. Plants with similar implementation of pores are also known from the African continent (Zimbabwe) as those of Mitchell 375 (BM); see Eddy (1985), Fig. 28 E. Since the distin-guishing features given by Crum (1992a) do not allow a sure identification, Sphagnum bourbonense will be considered as a synonym of S. capense.Habitat: moist to wet, acid, mostly anorganic sub-strates, on rocks, near springsDistribution: Afr.2 (Malawi, Zimbabwe), Afr.3, Afr.4 (Madagascar, Réunion)

Sphagnum sect. Acrosphagnum 207

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120. S. ceylonicum Mitt. ex Warnst. in Hedwigia 29: 195. 1890 (Pl. 93)Syn.: S. kerstenii Hampe in Warnst. 1911, S. keniae Dixon 1918, S. vandenbroeckii Nav. 1922, S. ugandense J. Taylor & A. Thompson 1955Habitus: rather small to robust plants; colour: green, grey green, pale brown, yellow-brown to orange brown; stem: cortex two- to three-layered, rarely four-layered, cells of outer layer often wider, on the external surface with 0–2 pores at the cells ends, wood cylinder yellowish, orange to dark brown; stem leaf: ovate-lin-gulate to triangular-lingulate with rounded to obtuse, dentate or somewhat fringed, occasionally cucullate apex, size variable, on Asian plants 1.3–1.6 mm, on African plants 1.8–2.6 mm; border: 2–4 rows of cells wide, not widened downwards; hyaline cells: mostly undivided, rarely some one time divided in the upper half of leaf, with fibrils down to the middle of leaf or to the leaf base, on ventral side with single pores in apical cell angles or with scattered small pseudopores in cell angles, on dorsal side with large (7–12 μm), round, mostly ringed, strung pores along the commis-sures or dispersed over the whole cell areas or numer-ous large membrane gaps, near leaf base pores mostly in cell angles; branch fascicle: with 4–6 (–7), rather weakly dimorphous, but different in length, occasion-ally 2 more robust branches spreading, the other pend-ing, cortex with often 2–3 retort cells in rows, branch leaves mostly not five-ranked, sometimes indistinctly five-ranked; branch leaf: ovate to ovate-lanceolate with more or less obtuse, narrowly truncated and den-tate leaf tip, (1.0–) 1.4–1.8 (–2.1) mm long; branch leaf pores: on ventral side pores lacking or scattered, weakly ringed pores in cell angles, occasionally with several pseudopores in cell angles, on dorsal side mostly with medium-sized (5–8 μm, Africa) or large (6–10 μm, Asia), elliptic, ringed pores in dense rows along the commissures and additionally with 1–6 large, round, ringed, centred pores; chlorophyll cells: invert triangular to trapezoid, ventral broadly exposed, dorsal enclosed or narrowly exposed; hyaline cells: ventral almost plane, dorsal convex, walls adjacent to chlorophyll cells smooth.Remarks: On basis of the leaf cross-section and the relatively short and wide hyaline cells Warnstorf (1911) assigns this species (written as S. ceylanicum) to S. sect. Acutifolia. Nevertheless, it fits better to the of-ten poly-pored S. sect. Acrosphagnum (cf. Eddy 1977). Eddy (1977, 1985) mentions different measurements for African and Asian plants but don’t separate them on species level.

Habitat: acid oligotrophic to mesotrophic bogs, on rocksDistribution: As.3 (Sri Lanka, ? South India), Afr.2 (Uganda, Kenya, Zaire), Afr.4 (Madagascar, Rèunion)

121. S. complanatum Flatberg & Whinam in J. Bryol. 33: 112. 2011 (Pl. 94)Habitus: small, short and densely branched, column-shaped plants with conical to roundish capitula; colour: reddish, light orange to almost whitish green, occasion-ally with pink tinge; stem: cortex mostly two-layered, cells of outer layer distinctly wider, on the external surface often with one large apical pores, sometimes without pores, wood cylinder pale to light orange; stem leaf: elliptic to ovate with involute, acute apex with of-ten 2–4 denticles, 2.0–2.6 mm long; border: 1–3 rows of cells wide, not widened downwards, sometimes lacking at the apex; hyaline cells: with fibrils almost down to the leaf base, undivided, on ventral side with up to 16 ringed pores and/or pseudopores along the commissures, on dorsal side with numerous round to elliptic, mostly ringed commissural pores, occasion-ally pores also apart from the commissures (subcom-missural) or centred and often slightly enlarged, near the leaf base pores lacking; branch fascicle: with 1–3, thereof 2 stronger and about 5 mm long and 1 weaker and about 3.5 mm long, all obliquely emergent or the weaker branch rather pending, the stronger branches with flattened leaf arrangement, the weaker branch rather indistinctly flattened, cortex with often two re-tort cells, one after another, branch leaves not or only indistinctly five-ranked; branch leaf: ovate-lanceolate with involute, more or less dentate leaf tip, straight or slightly secund, 1.3–1.9 mm long; branch leaf pores: implementation of pores similar to stem leaf, but on dorsal side subcommissural and centred pores more frequent; chlorophyll cells: elliptic, centred, on both sides with thickened cell walls exposed or apparently enclosed; hyaline cells: on both sides slightly convex, walls adjacent to chlorophyll cells smooth.Habitat: in moist to wet moss lawns, on peaty soil, often together with Racomitrium lanuginosumDistribution: Afr.3 (Île Amsterdam)

122. S. davidii Warnst., Allg. Bot. Z. Syst. 11: 99. 1905 (Pl. 95–96)Syn.: S. chevalieri Warnst. 1911, S. afro-crassicladum Dixon et Sherrin 1938Habitus: rather small to medium-sized, sometimes robust plants; colour: pale or dirty brownish to yel-low-brown, occasionally green; stem: cortex two- to three-layered, cells of outer layer distinctly wider, on

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sthe external surface partly with one large, apical pore, wood cylinder yellowish, yellow-brown, red to dark reddish brown; stem leaf: triangular-lingulate to lin-gulate, rounded or obtuse, occasionally cucullate apex, 1.1–1.6 (Warnstorf 1911) resp. 1.6–2.6 mm long (Eddy 1985); border: 2–4 rows of cells wide, not widened downwards; hyaline cells: mostly undivided, occasion-ally in the upper half one to several times divided, with fibrils in the upper half of leaf or down to the leaf base, on ventral side pores almost lacking, on dorsal side with many small to medium-sized, ringed or unringed, densely strung commissural pores; branch fascicle: with 4–7, not or weakly dimorphous, 2–3 spreading, cortex with often two retort cells, one after another, sometimes neighbouring cortex cells also porous, branch leaves more or less indistinctly five-ranked; branch leaf: ovate (in the middle of branch) to ovate-lanceolate (in the proximal part of branch) with nar-rowly truncated leaf tip, often with few denticles, of-ten slightly asymmetric, 1.0–1.5 mm long; branch leaf pores: on ventral side variable, often pores resp. pseu-dopores lacking or few, occasionally poly-porous, on dorsal side with numerous small to medium-sized (4–6 μm), ringed pores in rows along the commissures, with or without 1–6 additional centred pores; chloro-phyll cells: narrowly elliptic to elliptic-rectangular, cen-tred, on both sides with thickened cell walls exposed; hyaline cells: ventral convex to slightly convex, dorsal convex, walls adjacent to chlorophyll cells smooth.Habitat: acid oligotrophic to mesotrophic mires, Eri-caceae-dominated vegetation, on wet rocksDistribution: Afr.2, Afr.4

123. S. ericetorum Brid., Muscol. recent. Suppl. 1: 19. 1806 (Pl. 97)Habitus: small plants; colour: green, yellowish to brown; stem: cortex two- to three-layered, wood cyl-inder yellowish; stem leaf: triangular-lingulate to al-most lingulate with an obtuse, shorter or somewhat longer, cucullate apex, due to involute margins appar-ently more acute, 1.7–2.5 mm long; border: very nar-row, not widened downwards; hyaline cells: undivided, with fibrils throughout the whole leaf, on ventral side with large, round, mostly unringed, commissural pores in rows, getting larger downwards up to cell width, on dorsal side only single pores in cell angles; branch fas-cicle: with 4–6, hardly to weakly dimorphous, in rather dimorphous branch fascicles often 2 branches spread-ing, cortex with often two retort cells, one after anoth-er; branch leaf: ovate with narrowly truncated, dentate leaf tip, strongly concave, about 1.4 mm long; branch leaf pores: on ventral side with numerous, rather large

(6–10 μm), round, ringed commissural pores and part-ly similar centred pores, on dorsal side pores lacking or with few pores in cell angles, more numerous near leaf margins and leaf tip, often also with several pseu-dopores; chlorophyll cells: ovate to ovate-triangular, ventral narrowly exposed or scarcely enclosed, dorsal mostly with thickened cell wall broadly exposed, only sometimes also enclosed; hyaline cells: ventral convex, dorsal almost plane, walls adjacent to chlorophyll cells smooth.Remarks: Warnstorf (1911) assigned this species to S. sect. Cuspidata. Eddy (1985) discussed similarities with species of the Acrosphagnum group, especially with S. pycnocladulum, and mentioned that it could be-long to that species.Habitat: wet sites in heath vegetationDistribution: Afr.4 (Madagascar, Rèunion)

124. S. islei Warnst. in Hedwigia 30: 14. 1891 (Pl. 98)Habitus: small plants with flattened to slightly convex capitula; colour: patchy orange to pale yellow-brown; stem: cortex one- to two-layered, on the external sur-face without pores or with one apical pore, wood cylin-der pale to yellowish; stem leaf: ovate, ovate-lingulate to ovate-triangular with narrowly rounded, often invo-lute and apparently acute apex, sometimes very slightly falcate, 1.3–1.9 mm long; border: narrow, not widened downwards; hyaline cells: undivided, with fibrils in the upper one or two thirds, occasionally down to the leaf base, on ventral side mostly without pores, on dorsal side in the upper half (or two thirds) with numerous small, ringed commissural pores, getting downwards larger, weakly ringed and less numerous; branch fas-cicle: with 1–3, thereof often 2 more robust spreading, relatively short, about 5 mm long, especially spreading branches more or less distinctly five-ranked; branch leaf: ovate to ovate-lanceolate with narrow to pointed leaf tip, often slightly falcate, 0.9–1.3 mm long; branch leaf pores: on ventral side pores almost lacking, on dor-sal side in the upper half with numerous small, ringed commissural pores; chlorophyll cells: elliptic, on both sides with thickened cell walls narrowly exposed or on dorsal side relatively broadly exposed; hyaline cells: on both sides slightly convex, walls adjacent to chloro-phyll cells smooth.Remarks: Sphagnum islei is assigned as a synonym to S. capense by Eddy (1985), but revised as a separate species by Flatberg et al. (2011). The main distinguish-ing features are the not broadly rounded stem leaves and the rather lanceolate, not truncated branch leaves. The descriptions differ in some details. Both, Warnstorf (1891, 1911) and Flatberg et al. (2011) show centred,

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more or less enclosed chlorophyll cells in branch leaf cross-sections, whereas Eddy (1985) mentions on dor-sal side rather broadly exposed chlorophyll cells on ba-sis of the type specimen.Habitat: on high level at the Caldera Plateau near the Lac BleuDistribution: Afr.3 (Île Amsterdam)

125. S. novo-caledoniae Paris & Warnst. apud Warnst. in Engl., Nat. Pflanzenfam.I, 3: 297. 1911 (Pl. 99)Syn.: S. novae-caledoniae Paris & Warnst. in Broth. 1910 nom. nud., Flatbergium novo-caledoniae Devos et al. 2016 nom. nud.Habitus: very small plants with small and rather indis-tinct capitula; colour: above grey green, below ochra-ceous; stem: cortex one- to two-layered, on the external surface without pores, wood cylinder above pale, below dark brown to almost black; stem leaf: elongated ovate-lanceolate with rounded-truncated, dentate or some-what fringed apex, more or less concave, 1.4–1.8 mm long; border: very narrow or almost dwindling; hyaline cells: in the lower half of leaf occasionally divided, always without fibrils, on ventral side with scattered small pores in cell angles, in the upper half with mem-brane gaps, on dorsal side densely multi-porous with differently large, roundish to elliptic, unringed pores; branch fascicle: single or with 2 (–3), very short (4–5 mm long), cortex with retort cells; branch leaf: elon-gated ovate with rounded, fringed to dentate leaf tip, 0.7–1.1 mm long; hyaline cells without fibrils; branch leaf pores: on ventral side with small single pores in cell angles, on dorsal side densely multi-porous with differ-ently large, roundish to elliptic or irregular, unringed pores; chlorophyll cells: invert triangular to trapezoid, ventral broadly exposed, dorsal narrowly exposed or enclosed; hyaline cells: ventral almost plane, dorsal convex, walls adjacent to chlorophyll cells smooth.Remarks: The placement of this unconventional spe-cies to a certain section is not clarified satifactorily. An assignment to S. sect. Acrosphagnum would be sup-ported by the relatively broad and multiporose hyaline cells and the enlarged outer cells of the stem cortex. A tendency of reduction of fibrils in the branch leaf hya-line cells is also visible in multiporose forms of other species like S. capense.Devos et al. (2016) use the name Flatbergium no-vo-caledoniae (also written as: ʻFlatbergium novo-caledoniaeʼ) with reference to J. Shaw, unpublished. Without any other explanation, the name Flatbergium novo-caledoniae (Paris & Warnst.) Devos et al. has to be considered as a nomen nudum.

In recent time, Müller (2012) describes the sporophyte of S. novo-caledoniae which was unknown before. The pseudopodium is very short; the capsule is sitting between perichaetial leaves on an elongated and laxly foliated branch. Pseudostomata are apparently absent from the capsule surface.Habitat: on moist stones in streams (Iwatsuki 1986), silty river bank along small river (TRH B-9337)Distribution: Oc. (New Caledonia)

126. S. perrieri Ther. in Recueil. Publ. Soc. Natl. Havraise Études Diverses 39: 112. 1922 (Pl. 100)Habitus: small, densely branched plants with hemi-spherical capitula; colour: pale; stem: cortex one- to three-layered, wood cylinder pale; stem leaf: triangu-lar-lingulate, lingulate to ovate-lingulate with cucullate apex, 1.3–1.4 mm long; border: in the middle of leaf 3–5 rows of cells wide, not or hardly widened down-wards; hyaline cells: undivided, with fibrils in the up-per four fifths of the leaf, on ventral side few round, 7–8 μm wide, unringed pores along the commissures and in cell angles, getting more numerous near leaf margins, on dorsal side with numerous round to irregular, 10–15 μm wide commissural pores, in broad hyaline cells also with centred pores; branch fascicle: with 4, mod-erately dimorphous, rather short, (2–) 3–6 mm long, leaves especially on spreading branches five-ranked; branch leaf: lanceolate, ovate-lanceolate to ovate with short, obtuse leaf tip, 0.7–1.0 mm long; branch leaf pores: on ventral side pores completely or almost lack-ing, occasionally with scattered round, unringed pores in cell angles, on dorsal side with numerous round to irregular, 10–15 μm wide commissural pores, in broad hyaline cells also with centred pores; chlorophyll cells: rectangular, narrowly barrel-shaped to slightly invert trapezoid, centred, on both sides with thickened cell walls exposed, often somewhat broader on ventral side; hyaline cells: on both sides slightly convex, walls adja-cent to chlorophyll cells smooth.Remarks: Sphagnum perrieri is assigned to S. sect. Sub- secunda subsect. Subsecunda in Michaelis (2011). Af-ter examination of herbarium specimens (PC 0085849), this species will allocated to S. sect. Acrosphagnum because of the relatively short and broad branch leaf hyaline cells with large pores and the enlarged outer cells of stem cortex.Habitat: not mentionedDistribution: Afr.4 (Madagascar)

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Plate 163: Sphagnum rio-negrense: a: habitus, b: stem leaves, c: stem leaf cells ventral, d: stem leaf cells dorsal, e: stem leaf cross-section; Sphagnum rotundifolium: f: stem leaves, g: branch leaves, h: branch leaf cross-sections (above in water, be-low in diluted sulphuric acid); Sphagnum subovalifolium: i: stem leaves, j: branch leaves, k: branch leaf cross-sections (like in h); Sphagnum subrufescens: l: stem leaves, m: branch leaf, n: branch leaf cross-section; Sphagnum turgescens: o: habi-tus, p: stem leaves, q: branch leaf, r: lower branch leaf cells dorsal, s: branch leaf cross-section.

318 Species descriptions / plate 163

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Ambuchanania 10, 19, 27leucobryoides 20, 29, 353, 397, Pl. 194

Ambuchananiaceae 15, 29Andreaea 10, 20Bryum 23Calliergon

trifarium 233Cryphea

heteromalla 12Drosera

rotundifolia 11Eosphagnum 15, 29

inretortum 15, 20, 73rigescens 73

Flatbergiaceae 15, 28Flatbergium 28

novo-caledoniae 210sericeum 15, 20, 28

Fissidens 23Isocladus 12, 14

macrophyllus 14, 331Junjagia

glottophylla 17Leucobryum

falcatum 12Muscus

terrestris vulgaris 12Polytrichum 11Protosphagnum

nervatum 17Racomitrium

lanuginosum 208Scorpidium 233Sphagnophyllites

triassicus 17, 19Sphagnum

sect. Acisphagnum 14, 31sect. Acocosphagnum 14, 15, 21, 28, 37, 67, 73sect. Acrosphagnum 21, 27, 28, 31, 40, 44, 65, 67, 68,

71, 155, 207, 208, 209, 210sect. Acutifolia 7, 12, 13, 14, 15, 18, 19, 20, 21, 27, 29,

30, 33, 38, 44, 48, 56, 65, 66, 67, 68, 69, 70, 71, 84, 132, 136, 155, 208, 231

sect. Anacamptosphagnum 30sect. Cavifolia 30sect. Comatosphagnum 14, 30sect. Cuculliformes 29sect. Cuspidata 7, 9, 10, 12, 13, 14, 15, 18, 20, 21, 28,

31, 32, 35, 36, 42, 46, 50, 52, 63, 66, 67, 68, 69, 70, 71, 209, 332, 338, 340, 352

sect. Cymbifolia 12, 13, 14, 29sect. Eusphagnum 15sect. Hemitheca 21, 27, 32, 37, 46, 54, 65, 66, 68, 69,

70, 331sect. Inretorta 21, 27, 28, 29, 54, 69, 70, 73, 231sect. Insulosa 15, 21, 27, 29, 33, 38, 48, 66, 67, 68, 91

9. Index

sect. Inophloea 15sect. Inundata 30sect. Isocladus 15, 21, 28, 31, 52, 69sect. Lapazensis 28, 29sect. Litophloea 15sect. Malacosphagnum 14, 29sect. Mollia 14, 136sect. Mollusca 12, 14, 21, 27, 32, 37, 43, 46, 54, 65, 66,

67, 68, 69, 70, 71, 352sect. Mucronata 15, 31sect. Palustria 13, 29sect. Platysphagnum 14, 29sect. Polyclada 9, 15, 21, 27, 30, 35, 40, 51, 66, 67, 69,

157sect. Pycnoclada 30sect. Pycnosphagnum 14, 30sect. Rigida 7, 9, 12, 14, 15, 20, 21, 27, 29, 33, 38, 44,

47, 56, 65, 66, 67, 68, 69, 70, 71, 89sect. Sericea 28sect. Sphagnum 7, 9, 10, 13, 15, 17, 19, 20, 21, 27, 29,

33, 36, 43, 46, 54, 65, 66, 67, 68, 69, 70, 71, 73, 74, 84, 231

sect. Squarrosa 7, 9, 14, 15, 20, 21, 27, 30, 35, 40, 44, 50, 65, 66, 67, 68, 69, 71, 156

sect. Subsecunda 7, 10, 12, 13, 14, 15, 19, 20, 21, 27, 28, 29, 30, 31, 35, 40, 45, 51, 58, 65, 67, 68, 69, 70, 71, 84, 143, 210, 211, 225, 231, 331, 332, 338, 340, 342, 352

sect. Truncata 14, 29subgen. Acutifolia 15subgen. Cuspidata 15[subgen.] Heterophylla 12, 14[subgen.] Homophylla 12, 13, 14, 15[subgen.] Inophloea 13[subgen.] Isocladus 13, 15[subgen.] Litophloea 13[subgen.] Rigida 13, 15[subgen.] Sphagnum 13, 15subgen. Squarrosa 15subgen. Subsecunda 15subsect. Acrosphagnum 19, 31, 211subsect. Acutifolia 15, 30, 135subsect. Cuspidata 15, 31subsect. Cymbifolia 15subsect. Fimbriata 30, 132, 141subsect. Mucronata 15, 31subsect. Polyclada 15subsect. Rigida 15subsect. Sericea 15subsect. Squarrosa 15subsect. Subsecunda 15, 31, 210subsect. Truncata 15aciphyllum 57, 69, 70, 141, 142, 143, 178, Pl. 61aconiense 156, 157aculeatum 211acutifolioides 40, 67, 152, 194, Pl. 77

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426 Index

acutifolium 12, 13, 14, 15, 144 ssp. iridum 142 ssp. luridum 153 var. aquaticum 153 var. borbonicum 141 var. decipiens 139 var. elegans 138, 144 var. fallax 133, 139 var. fl agelliforme 139 var. fl avicomans 154 var. fl avicaule 149 var. fuscescens 137 var. fuscoluteum 137 var. fuscum 137 var. gerstenbergeri 150 var. gracile 142 var. graefi i 142 var. intermedium 142 var. laxum 153 var. luridum 153 var. meridense 147 var. pallens 130 var. plumosum 153 var. polyphyllum 139 var. purpureum 153 var. quinquefarium 142, 149 var. roseum 139 var. robustum 139 var. schillerianum 153 var. schimperi 144 var. schliephackeanum 144 var. silesiacum 150 var. squarrulosum 153 var. strictiforme 139 var. subtile 144 var. tenellum 138acutiforme var. auriculatum 139 var. elegans 139 var. fallax 133 var. fuscum 137 var. graeffi i 150 var. robustum 139 var. rubellum 138 var. tenellum 138, 142, 150acutirameum 62, 70, 211, 227, 260, Pl. 105acutum 332 var. hakusanense 332aequalipunctatum 61, 70, 212, 260, Pl. 105aequifolium 243, 327aequiporosum 58, 69, 70, 143, 212affi ne 22, 33, 43, 47, 54, 66, 68, 69, 70, 74, 75, 94, 238,

Pl. 3 var. affi ne 74, 75, 94 var. fl agellare 74, 75, 94africanum 45, 68, 227, 233, 236, 306, 340, Pl. 151afro-crassicladum 208alabamae 220 var. humile 220

alaskanum 338alaskense 37, 47, 66, 67, 68, 76, 99, Pl. 8albescens 347albicans 339, 369alegrense 47, 54, 69, 70, 76, 100, Pl. 9algentryi 87allionii 87aloysii-sabaudiae 344alpinum 144, 344amazonense 58, 69, 143, 179, Pl. 62amazonicum 55, 70, 82, 112, Pl. 21ambiguum 90amblyphyllum 347americanum 138, 154amoenoides 15, 55, 70, 82, 112, Pl. 21amoenum 87andersonianum 15, 16, 138, 139, 144andinum 85andrusii 51, 69, 212, 261, Pl. 106angermanicum 34, 49, 66, 68, 136, 154, 198, Pl. 81angolense 233angustifolium 22, 25, 36, 43, 54, 66, 67, 69, 342, 343,

346, 378, Pl. 200angustifrons 244angusti-limbatum 339, 371anisoporum 139annulatum 22, 36, 43, 53, 66, 67, 69, 334, 337, 359,

Pl. 181 var. porosum 334, 336antarcticum 87antarense 41, 67, 212, 261, Pl. 106antillarum 87, 144, 146antioquiense 63, 69, 212, 262, Pl. 107aongstroemii 13, 14, 15, 22, 27, 29, 33, 38, 48, 66, 67,

68, 91, 131, 136, Pl. 40apiculatum 346apollinairei 151apopenneysii 344, 345aquaticum 344aquatile 215aracense 153arbogastii 87arboreum 85arcticum 33, 38, 48, 66, 132, 138, 158, Pl. 41armoricum 218artariae 235aschenbachianum 144assamicum 81atlanticum 351atroligneum 15, 87attenuatum 87aureum 47, 69, 82, 113, Pl. 22auriculatum 14, 215, 216, 238, 338 var. inundatum 218austinii 14, 15, 22, 33, 47, 66, 68, 74, 75, 76, 95, Pl. 4australe 10, 38, 44, 65, 67, 68, 71, 83, 89, 127, Pl. 36austro-americanum 57, 59, 70, 143, 145, 179, Pl. 62austro-molle 207azuayense 57, 69, 155, 200, Pl. 83

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bahiense 87 var. sincorae 76, 77bakeri 218, 220balfourianum 87balslevii 57, 69, 136, 166, Pl. 49balticum 22, 36, 42, 52, 64, 66, 67, 69, 70, 334, 335,

337, 346, 360, Pl. 182 var. ruppinense 334barclayae 64, 70, 335, 361, Pl. 183bartlettianum 16, 138, 139, 144 var. roseum 144bartlettii 77batumense 215bavaricum 218beccarii 87belli-imbricatum 85beothuk 22, 34, 48, 66, 68, 136, 167, Pl. 50bergianum 140beringiense 51, 69, 234, 307, 308, Pl. 152, 153bernieri 344bescherellei 90bessonii 339beyrichianum 211, 256, 257bicolor 85biforme 80, 87billbuckii 15, 77, 78boasii 157bocainense 59, 70, 230, 299, Pl. 144bohemicum 344bolanderi 132boliviae 60, 69, 70, 234, 309, Pl. 154boomii 62, 70, 213, 262, Pl. 107borbonicum 141bordasii 226, 227, 292boreale 334borneoense 78bostonense 218, 220bourbonense 207, 245, 246, 247boyacanum 65, 70, 343, 379, Pl. 201brachybolax 80, 87brachycaulon 235brachycladum 80, 87brasiliense 54, 70, 77, 78, 79, 103, Pl. 12breedlovei 49, 69, 136, 166, Pl. 49brevicaule 87brevifolium 346, 347brevirameum 54, 70, 77, 78, 79, 104, Pl. 13brotherusii 335brunnescens 144buckianum 15, 55, 70, 83, 113, Pl. 22bushii 218, 220caldense 231 var. scorpidioides 225caldensi-recurvum 347callichroum 225calymmatophyllum 40, 67, 213, 226, 236, 263, 264,

Pl. 108, 109campbellii 71, 89, 90campellianum 89

campicolum 153camusii 215capense 31, 45, 68, 207, 209, 245, 246, 247, 340, Pl. 90,

91, 92capillaceum 144 var. tenellum 138 var. tenerum 140capillifolioides 144capillifolium 12, 13, 21, 22, 30, 35, 39, 40, 44, 50, 58,

66, 67, 68, 69, 70, 132, 136, 139, 141, 143, 144, 145, 147, 180, Pl. 63

var. tenellum 138, 144 var. tenerum 140 var. viride 144cardotii 339carneum 87carolinianum 51, 69, 234, 305, Pl. 150cavernulosum 46, 68, 230, 231, 300, Pl. 145cavifolium 15, 225 var. laricinum 336centrale 22, 33, 47, 66, 67, 68, 83, 114, Pl. 23ceylonicum 40, 45, 67, 68, 155, 208, 248, Pl. 93chevalieri 208, 250, 251chi-chiense 61, 63, 70, 213, 265, Pl. 110 var. uvidulum 213, 265chilense 132chinense 12, 347chlorocephalum 80clandestinum 12cleefi i 62, 69, 213, 266, Pl. 111cochlearifolium 220, 238columniforme 56, 70, 79, 106, Pl. 15commutatum 237comosum 233, 237compactum 12, 13, 14, 21, 22, 29, 33, 38, 44, 47, 56, 65,

66, 67, 68, 69, 70, 71, 90, 128, 136, 236, Pl. 37 var. expositum 91 var. ovatum 89 var. ramulosum 135 var. rigidum 90complanatum 45, 68, 208, 249, Pl. 94concinnum 33, 48, 66, 68, 132, 135, 159, Pl. 42condensatum 12, 68, 90, 91, 129, Pl. 38confertum 89confl atum 61, 70, 214, 268, Pl. 113congoanum 339, 371connectens 42, 67, 335, 361, Pl. 183contortulum 63, 70, 219, 352, 394, Pl. 216contortum 13, 14, 21, 22, 35, 41, 51, 66, 67, 69, 234,

310, Pl. 155 var. laricinum 234 var. obesum 215 var. platyphyllum 238controversum 336convolutum 226cordemoyi 141, 175cordifolium 218, 220cornutum 215coronatum 226

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coryphaeum 147, 186costae 27, 56, 70, 135, 164, Pl. 47 var. confertorameum 135, 164 var. costae 135, 164 var. seriatum 135, 164costaricense 147crassicladum 215crassisetum 80, 156crassum 85cribriforme 61, 70, 228, 266, Pl. 111cribrosum 52, 69, 244, 329, 331, Pl. 173crispatum 336, 344, 345crispum 51, 69, 214, 267, Pl. 112cristatum 65, 71, 83, 115, Pl. 24cruegeri 144crumii 63, 70, 214, 268, Pl. 113cubile 80cucullatum 61, 70, 214, 267, Pl. 112cuculliforme 29, 55, 69, 70, 83, 84, 116, Pl. 25cundinamarcanum 55, 69, 84, 116, Pl. 25cupressiforme 215curicuriariense 59, 70, 231, 299, Pl. 144curvatulum 65, 70, 343, 380, Pl. 202 var. curvatulum 343, 380 var. steereanum 343, 380curvifolium 235cuspidatifolium 344cuspidatiforme 344cuspidatulum 42, 46, 67, 68, 332, 347, 350, 354, Pl. 176 var. fi brosum 332 var. fuscescens 332 var. malaccense 332 var. trengganuense 344cuspidatum 12, 13, 14, 15, 21, 22, 31, 36, 43, 46, 53, 65,

66, 67, 68, 69, 70, 71, 334, 336, 340, 344, 345, 351, 352, 380, 381, Pl. 203, 204, 205

fulvum 332 ssp. cuspidatum 344, 345, Pl. 203, 204 ssp. subrecurvum 71, 344, 345, 382, Pl. 205 var. brevifolium 346 var. cuspidatum 344, 345 var. defl exum 337 var. densum 332 var. dusenii 338 var. falcatum 66 var. fallax 346 var. fl accidifolium 383 var. latetruncatum 226 var. majus 337 var. miquelonense 351 var. mollisimum 334 var. plumosum 344, 345 var. roellii 334 var. serrulatum 42, 53, 63, 66, 341, 344, 345, 348,

381, Pl. 180 var. speciosum 333 var. subrecurvum 383 var. torreyi 351cyclocladum 61, 70, 214, 215, 269, Pl. 114

cyclophyllum 51, 59, 69, 70, 231, 301, Pl. 146cymbifolioides 80, 237cymbifolium 12, 13, 14, 80, 81 ssp. austinii 74 ssp. papillosum 78 var. compactum 85, 90 var. congestum 85 var. cordifolium 91 var. ludovicianum 87 var. medium 85 var. papillosum 78 var. paradisii 85 var. patens 147 var. purpurascens 85 var. squarrosum 156 var. tenellum 352 var. turgidum 231cymbophylloides 83cymbophyllum 83dasyphyllum 215, 217davidii 31, 45, 68, 208, 250, 251, Pl. 95, 96decipiens 83, 238defl exum 80degenerans 74delamboyense 61, 62, 70, 215, 270, Pl. 115densicaule 89densirameum 221densum 142denticulatum 14, 22, 25, 35, 40, 41, 45, 66, 68, 215,

216, 220, 225, 271, Pl. 116derrumbense 80, 87diblastoides 57, 69, 143, 145, 181, Pl. 64diblastum 142dicladum 38, 67, 155, 185, Pl. 68dielsianum 83dimorphophyllum 56, 70, 79, 106, Pl. 15discrepans 87dissimile 64, 70, 336, 345, 379, Pl. 201divinum 85, 86divisum 62, 70, 216, 272, Pl. 117domingense 91dominii 66, 71, 345, 379, Pl. 201d’Orbignyanum 85drepanocladum 335drouhardii 87drummondii 231dubiosum 237dubium 347dusenii 338 var. majus 338dusenioides 64, 70, 335, 336, 361, Pl. 183earlei 87eatonii 140ecuadorense 147efi brillosum 21, 28, 40, 67, 216, 272, Pl. 117ehyalinum 28, 63, 70, 336, 345, 346, 384, Pl. 206elegans 335elenkini 342, 343ellipticum 241

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engelii 64, 70, 346, 385, Pl. 207ericetorum 45, 68, 89, 141, 209, 252, Pl. 97erosum 89erythrocalyx 78, 87 var. papillosum 78eschowense 226evansii 140exile 59, 70, 216, 273, Pl. 118exquisitum 59, 70, 231, 302, Pl. 147falcatum 66falcatulum 10, 42, 65, 66, 67, 70, 71, 335, 336, 345,

362, Pl. 184falcifolium 235falcirameum 89fallax 22, 25, 36, 43, 53, 66, 67, 69, 343, 346, 347, 348,

350, 386, Pl. 208 var. fl exuosum 347faxonii 344feae 332fi mbriatum 11, 12, 13, 14, 15, 22, 30, 33, 38, 44, 48, 56,

66, 67, 68, 69, 70, 132, 135, 160, Pl. 43 ssp. concinnum 132 var. concinnum 132 var. strictum 133fi tzgeraldii 52, 64, 69, 70, 336, 345, 363, Pl. 185fl accidifolium 344fl accidum 61, 70, 217, 273, Pl. 118fl accirameum 243fl avicans 218, 220fl avicaule 57, 69, 145, 181, Pl. 64fl avicomans 49, 69, 138, 154, 199, Pl. 82fl avirameum 243fl avum 145fl eischeri 87fl exuosum 22, 37, 42, 43, 46, 54, 64, 66, 67, 68, 69, 70,

346, 347, 350, 387, Pl. 209 var. fallax 346 var. recurvum 347 var. tenue 342fl oridanum 244fl uctuans 226fl uitans 218, 220fontanum 60, 70, 235, 308, Pl. 153frahmii 85, 86franconiae 220fraudulentum 58, 70, 137, 168, Pl. 51fulvum 332funkiae 60, 69, 235, 311, Pl. 156fuscovinosum 66, 71, 217, 233, 274, Pl. 119fuscum 15, 22, 34, 39, 48, 66, 67, 69, 136, 137, 139,

169, Pl. 52 var. schimperi 138gabonense 344galipense 147garberi 91garysmithii 60, 70, 228, 295, Pl. 140gedeanum 152geheebi 87georgianum 331

geraisense 58, 70, 217, 275, Pl. 120girgensohnii 14, 15, 21, 22, 30, 34, 38, 44, 48, 66, 67,

68, 69, 133, 134, 135, 154, 161, Pl. 44 var. majus 139 var. roseum 139glaucovirens 87glaucum 80globicephalum 224globicomosum 85godmanii 133goetzeanum 211, 256, 257gomezii 49, 69, 145, 146, 182, Pl. 65gordjagini 139gracile 139, 144, 147, 344gracilescens 59, 70, 244, 324, Pl. 169 var. laxifolium 244grandifolium 87grandirete 85grasslii 51, 69, 228, 295, Pl. 140gravetii 215griffi thianum 87griseum 59, 70, 235, 308, Pl. 153grossum 85guadalupense 87guanabarae 59, 70, 232, 302, Pl. 147guatemalense 89guwassanense 41, 67, 226, 236, 310, Pl. 155guyonii 87hahnianum 85hakkodense 78hamiltonii 15, 85hampeanum 66harleyi 54, 70, 84, 117, Pl. 26harperi 87hartmannii 91hegewaldii 60, 69, 229, 296, Pl. 141helenicum 45, 68, 236, 312, Pl. 157helleri 344helmsii 89helveticum 90hendocinum see mendocinumhenryense 37, 47, 67, 68, 77, 101, 102, Pl. 10, 11hercynicum 218herminieri 75hertelianum 15, 56, 69, 79, 107, Pl. 16herteri 62, 70, 217, 275, Pl. 120heterophyllum 80, 87hildebrandtii 211holleanum 14, 73holtii 215, 218holttumii 87homocladum 237homophyllum 60, 70, 229, 295, Pl. 141hookeri 133huilense 88humbertii 207humidulum Besch. in C. Müll. 1900 nom. nud.humile 91, 135huntii 87

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husnotii 87hyalinum 156hymenophyllophilum 149hypnoides 226, 344ikongense 344illecebrum 331imbricatum 18, 37, 66, 67, 74, 75, 97, Pl. 6 ssp. affi ne 74 ssp. austinii 74 ssp. austinii var. arcticum 76 var. fuscescens 74immersum 78, 90imperforatum 15, 55, 69, 70, 84, 117, Pl. 26incertum 38, 39, 67, 146, 182, Pl. 65incommodum 59, 69, 232, 303, Pl. 148incundum 48, 69, 137, 138, 170, Pl. 53inexspectatum 41, 51, 67, 69, 226, 242, 321, Pl. 166inretortum 28, 29, 73, 74insulatum 91insulosum 91intermedium 83, 144, 344 var. compactum 90 var. pulchrum 349inundatum 14, 21, 22, 35, 41, 45, 46, 51, 66, 67, 68, 69,

217, 218, 225, 276, Pl. 121iridans 12irritans 335irwinii 54, 70, 78, 104, Pl. 13islei 45, 68, 209, 253, Pl. 98isoloma 133isophyllum 238isoviitae 15, 346, 347itabense 63, 70, 218, 276, Pl. 121itacolumitis 77, 78itatiaiae 147, 151japonicum 80 var. philippinense 87javanense 12javanicum 344javense 12jensenii 22, 36, 43, 53, 66, 67, 334, 336, 337, 364, Pl. 186 var. annulatum 334juliforme 63, 69, 218, 277, Pl. 122junghuhnianum 39, 49, 50, 65, 67, 71, 151, 152, 195,

Pl. 78 var. junghuhnianum 195 var. pseudomolle 195kearneyi 351kegelianum 80, 87kelantanense 87kenaiense 334, 335keniae 208, 248kerstenii 208khasianum 41, 67, 219, 278, Pl. 123kihlmanii 333kiiense 152kinlayanum 225kinlayi 225kirkii 335

klinggraeffi i 80krylovi 347kurzeanum 344kurzianum 133kushiroense 41, 67, 219, 279, Pl. 124labradorense 135laceratum 56, 70, 133, 141, 158, Pl. 41lacteolum 90laegaardii 64, 70, 337, 365, Pl. 187lanceolatum 335lancifolium 335langloisii 218, 220lankesteri 52, 69, 337, 365, Pl. 187lapazense 19, 20, 21, 29, 54, 69, 73, 93, Pl. 2laricinum 14, 66, 234, 336, 349 var. platyphyllum 238late-limbatum 226laterale 144late-truncatum 235laticoma 237latifolium 80 var. compactum 90 var. condensatum 90 var. cordifolium 91 var. squarrosum 156laxifolium 14, 344 var. dusenii 338 var. majus 337 var. miquelonense 351 var. serrulatum 344laxirameum 57, 69,70, 146, 183, Pl. 66laxiramosum 61, 70, 219, 277, Pl. 122laxirete 90laxulum 63, 70, 220, 280, Pl. 125laxum 220lechleri 147lehmannii 344 var. robustum 347leionotum 83lenense 36, 42, 52, 66, 67, 69, 332, 335, Pl. 177leonii 59, 70, 232, 303, Pl. 148leptocladum 13leratianum 87lescurii 20, 51, 62, 69, 70, 216, 218, 220, 225, 234, 281,

Pl. 126lesueurii 146leucobryoides 353lewisii 57, 69, 146, 183, Pl. 66liesneri 61, 69, 220, 280, Pl. 125ligulatum 347limbatum 50, 69, 146, 148, 182, 244, Pl. 65limprichtii 338lindbergii 13, 14, 15, 22, 36, 42, 52, 64, 66, 67, 69, 70,

141, 332, 356, Pl. 178 ssp. lenense 332 var. microphyllum 332lindmanii 217linguaefolium 335lingulatum 226, 347

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livonicum 334lojense 62, 69, 221lonchocladum 89lonchophyllum 344longicomosum 63, 70, 215, 221, 274, Pl. 119longifolium 344, 347longistolo 55, 69, 70, 85, 118, Pl. 27loricatum 85louisianae 218, 220ludovicianum 87luetzelburgii 61, 69, 229, 297, Pl. 142luridum 153luzonense 40, 67, 221, 282, Pl. 127macrocephalum 89, 127macromolluscum 339, 372macrophyllum 12, 21, 31, 52, 69, 328, 329, 331, Pl. 173,

174 var. burinense 331 var. fl oridanum 244, 331 var. macrophyllum 331macroporum 87macro-rigidum 89madegassum 339maegdefraui 58, 69, 147, 184, Pl. 67magellanicum 12,14, 21, 22, 33, 37, 44, 55, 66, 67, 68,

69, 70, 82, 85, 86, 88, 119, 120, Pl. 28, 29 ssp. grandirete 44, 68, 85, 86, 120, Pl. 29magistri 58, 70, 221, 282, Pl. 127magnifolium 215majus 22, 36, 42, 53, 66, 67, 69, 337, 338, 366, Pl. 188malaccense 332mandonii 153, 154maori-compactum 83margaritaceum 85margaritae 50, 69, 147, 184, Pl. 68marginatum 226 var. convolutum 226marlothii 87mathieui 226, 292matogrossense 55, 70, 86, 118, Pl. 27mauritianum 226, 243maximum 83mcqueenii 334, 336medium 85, 86, 138mehneri 133mendocinum 53, 69, 336, 338, 367, Pl. 189meridense 50, 57, 58, 68, 69, 70, 147, 151, 186, 244,

Pl. 69mexicanum 91microcarpum 231microcephalum 83, 147, 244microcuspidatum 63, 70, 347, 385, Pl. 207microphyllum 132microporum 41, 67, 219, 222, 226, 283, Pl. 128 var. junsaiense 240mildbraedii 90minoense 67minutulum 60, 70, 78, 236, 311, Pl. 156miquelonense 351, 395

mirabile 60, 70, 237, 311, Pl. 156mirum 40, 50, 69, 156, 202, Pl. 85mississippiensis 53, 69, 348, 388, Pl. 210missouricum 218, 238miyabeanum 225, 226mobilense 217, 220mohrianum 336molle 15, 22, 27, 34, 38, 48, 56, 66, 68, 69, 70, 135,

136, 165, Pl. 48 var. limbatum 133, 138, 154 var. molluscoides 154molliculum 237, 335mollissimum 207molluscoides 135molluscum 13, 14, 15, 91, 352monocladum 344monzonense 85moorei 237moronum 57, 69, 148, 187, Pl. 70mosenii 153mossmannianum 237mougeotii 347mucronatum 211, 346muelleri 135mülleri 13, 14, 135, 136multifi brosum 37, 67, 80, 108, Pl. 17multiporosum 55, 70, 86, 121, Pl. 30nano-porosum 335nanum 144, 352natans 342naumannii 335nawaschinii 338neglectum 235, 238negrense 56, 69, 70, 80, 107, Pl. 16nemoreum 144, 145 var. angermanicum 154 var. luridum 153 var. subtile 144 var. tenerum 140nepalense 38, 67, 155, 200, Pl. 83nicholsii 218, 220nitidulum 44, 68, 138, 166, Pl. 49nitidum 140, 153noryungasense 61, 70, 222, 284, Pl. 129novae-caledoniae 210novae-seelandiae 237novo-caledoniae 9, 19, 20, 28, 65, 71, 210, 231, 254,

Pl. 99novo-fundlandicum 218, 220novo-guineense 42, 68, 333, 357, Pl. 179novo-zelandicum 31, 66, 71, 89, 225, 237, 313, Pl. 158obesum 215, 241obliquefi brosum 59, 70, 222, 284, Pl. 129oblongum 80, 156obovatum 243, 325, 326, 327obtusifolium 80, 90, 219 var. compactum 90 var. condensatum 90 var. minus 90

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obtusiusculum 141obtusum 22, 36, 42, 52, 66, 67, 69, 338, 368, Pl. 190 var. dusenii 337ochraceum 78, 157ocultifolium 144okamurae 222olafi i 34, 48, 66, 69, 138, 148, 188, Pl. 71oligodon 226, 292oligoporum 222orbignyanum 85oregonense 338orgaosense 87orientale 41, 51, 67, 68, 69, 242, 322, Pl. 167orlandense 217ornatum 15, 55, 69, 70, 87, 121, Pl. 30orthocladum 89otagoense 83ouropretense 87ovalifolium 59, 70, 237, 314, Pl. 159 var. japonicum 213ovatum 41, 67, 85, 238, 314, Pl. 159oxycladum 226 var. mauritianum 226oxyphyllum 57, 70, 149, 187, Pl. 70pachycladum 83pacifi cum 53, 69, 348, 388, Pl. 210pallens 38, 67, 133, 158, Pl. 41pallidum 243, 325, 326, 327palustre 12, 15, 20, 22, 29, 33, 38, 43, 47, 56, 65, 66,

67, 68, 69, 70, 71, 77, 79, 80, 109, 144, Pl. 18 ssp. intermedium 83 ssp. medium 85 ssp. papillosum 78 var. capillaceum 144 var. capillifolium 144 var. compactum 90 var. congestum 85 var. medium 85panduraefolium 207papillosum 22, 33, 37, 43, 46, 54, 66, 67, 68, 70, 77, 78,

79, 105, Pl. 14 var. intermedium 83 var. leve 79 var. plumosum 77pappeani 207pappeanum 29, 38, 44, 67, 68, 90, 91, 129, Pl. 38paraguense 85paranae 80, 87paranense 63, 70, 222, 285, Pl. 130parcoramosum 63, 70, 223, 285, Pl. 130parvifolium 342parvulum 144patagoniense 335patens 90, 147patulum 139,156paucifi brosum 87pauciporosum 87pauloense 87pellucidifolium 244

pendulirameum 58, 70, 149, 189, Pl. 72pentastichum 7, 349perfoliatum 16, 242perforatum 60, 70, 238, 315, Pl. 160 var. rotundifolium 228perichaetiale 14, 37, 43, 44, 47, 55, 65, 67, 68, 69, 70,

71, 77, 78, 86, 87, 88, 122, 123, Pl. 31, 31, 33 var. tabuleirense 55, 87, 124, Pl. 33permolle 347perrieri 45, 68, 210, 255, Pl. 100personatum 64, 70, 239, 336, 339, 373, Pl. 195peruvianum 87pilaiei 331planifolium 10, 46, 68, 339, 340, Pl. 191, 192, 193, 194 var. angustilimbatum 339, 371, Pl. 193 var. congoanum 339 var. planifolium 339, 340, 369, 370, Pl. 191, 192 var. rugegense 46, 339, 340, 372, Pl. 194platycladum 147, 148platyphylloides 41, 60, 67, 70, 238, 316, Pl. 161platyphyllum 14, 21, 22, 35, 42, 52, 66, 67, 69, 238,

317, Pl. 162plicatum 217, 220plumosum 344plumulosum 12, 153, 154 var. fl avicomans 154 var. gerstenbergeri 150 var. quinquefarium 150pluriporosum 59, 70, 242, 323, Pl. 168poasense 53, 69, 348, 349, 389, Pl. 211polyphyllum 346polyporum 225porosum 157, 336portoricense 37, 47, 54, 67, 68, 69, 70, 74, 75, 98, Pl. 7potieri 339praemorsum 90priceae 58, 69, 155, 185, Pl. 68procerum 89propinquum 336prostatum 331pseudo-acutifolium 151pseudobesum 215pseudocontortum 218pseudocuspidatum 347pseudo-cuspidatum 339pseudocymbifolium 38, 67, 81, 110, Pl. 19pseudomedium 83pseudomolle 152pseudomolluscum 344pseudopatulum 139pseudoplatyphyllum 238pseudoramulinum 61, 70, 223, 286, Pl. 131pseudorecurvum 347pseudo-recurvum 347pseudo-rigidum 85pseudorufescens 237 var. fl avescens 237 var. fuscorufescens 237 var. virescens 237

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pseudoserratum 344pseudosquarrosum 218, 220pseudoturgidum 215, 218pugionatum 211puiggari 80, 87pulchellum 144, 151pulchricoma 339, 349, 350pulchrum 22, 36, 43, 53, 66, 67, 69, 349, 390, Pl. 212pulvinatum 58, 70, 149, 189, Pl. 72pumilum 241pungens 218pungifolium 43, 67, 349, 391, Pl. 213purpuratum 58, 70, 141, 153, 196, Pl. 79purpureum 138, 141, 153pusillo-squarrosum 133pusillum 344pycnocladulum 31, 45, 68, 209, 211, 256, 257, Pl. 101,

102pycnocladum 157pylaesii 27, 32, 37, 46, 54, 65, 66, 68, 69, 70, 330, 331,

332, Pl. 175pylaiei 331quinquefarium 7, 17, 22, 35, 39, 49, 66, 67, 69, 149,

190, Pl. 73quinquefolium 150ramosissimum Lour. in Steud. 1824 nom. nud.ramulinum 59, 70, 223, 232, 303, Pl. 148reclinatum 15, 56, 70, 81, 111, Pl. 20recurvatum 339recurviforme 344recurvum 7, 13, 14, 46, 53, 64, 68, 69, 70, 71, 340, 346,

347, 349, 350, 392, Pl. 214 ssp. amblyphyllum 347 ssp. angustifolium 342 ssp. balticum 334 ssp. mucronatum 346 ssp. pulchrum 349 ssp. recurvum 350 var. amblyphyllum 347 var. brevifolium 347 var. fallax 338, 346 var. majus 347 var. mollissimum 334 var. mucronatum 25 var. obtusum 338 var. parvifolium 342 var. porosum 336 var. preuschoffi i 333 var. quinquefarium 349 var. recurvum 350 var. riparium 333 var. robustum 333 var. spectabile 333 var. tenue 342rehmannii 226reichardtii 44, 65, 68, 71, 156, 201, Pl. 84richardsianum 52, 69, 239, 315, Pl. 160rigescens 19, 21, 28, 29, 54, 69, 70, 73, 74, 93, Pl. 2rigidiforme 90

rigidulum 90rigidum 13, 14, 15, 90, 136 var. humile 135 var. teneriffae 90rio-negrense 59, 60, 70, 239, 318, Pl. 163riparioides 347riparium 15, 22, 36, 42, 52, 66, 67, 69, 333, 358, Pl. 180ripense 63, 70, 223, 286, Pl. 131rivulare 237, 338robinsonii 41, 67, 239, 319, Pl. 164robustum 12, 139rodriguezii 141rodwayi 336roellii 334roraimense 60, 70, 240roseotinctum 87roseum 151rothii 347rotundatum 59, 70, 90, 224, 287, Pl. 132rotundifolium 60, 70, 240, 318, Pl. 163rubellum 10, 13, 14, 15, 22, 34, 39, 44, 49, 57, 66, 67,

68, 69, 70, 138, 139, 144, 151, 171, Pl. 54rubiginosum 34, 38, 48, 66, 67, 69, 134, 162, Pl. 45rubrofl exuosum 52, 69, 333, 357, Pl. 179rufescens 215rufulum 332rugegense 68, 339, 340rugense 68ruppinense 334russowii 10, 12, 15, 21, 22, 34, 39, 49, 66, 67, 69, 134,

139, 172, Pl. 55rutenbergii 45, 68, 243, 325, 326, 327, Pl. 170, 171,

172ruwenzorense 339salvanii 243sancto-josephense 52, 64, 69, 70, 340, 374, Pl. 196sanguinale 55, 69, 70, 82, 88, 125, Pl. 34santanderense 62, 70, 224, 288, Pl. 133santosense 80, 87schiffneri 227schimperi 144 var. gracile 138 var. tenellum 138schliephackeanum 144schliephackei 344schofi eldii 49, 69, 150, 191, Pl. 74schultzii 344schwabeanum 62, 70, 224scorpidioides 63, 70, 225, 289, Pl. 134scortechinii 335scotiae 141sedoides 231, 331seemannii 344sehnemii 57, 70, 139, 168, Pl. 51semisquarrosum 156septatoporosum 59, 70, 243, 323, Pl. 168septatum 43, 67, 350, 388, Pl. 210sericeum 7, 12, 13, 14, 19, 20, 21, 28, 37, 67, 73, 92,

Pl. 1

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434 Index

seriolum 73serrae 347serratifolium 335serratum 341, 344 var. serrulatum 335serrulatum 335setchellii 335siamense 81simile 217, 220simplex 66, 71, 233, 304, Pl. 149simplicicaulis 56, 70, 81, 111, Pl. 20simplicissimum 215sincorae 76, 100sintenisii 87sipmanii 55, 69, 70, 88, 125, Pl. 34sitchense 133, 153, 154skyense 10, 34, 66, 150, 151, 192, Pl. 75slooveri 46, 68, 340, 375, Pl. 197smithianum 218, 220sociabile 147sonsonense 60, 70, 230, 298, Pl. 143sordidum 344sparsifolium 69, 90sparsum 49, 50, 57, 69, 70, 146, 151, 193, Pl. 76speciosum 333spectabile 14, 333spegazzinii 335sphaericum 66spinulosum 85splendens 28, 52, 69, 350, 389, Pl. 211squarrifolium 69squarrosiforme 152squarrosulum 157squarrosum 12, 13, 14, 15, 22, 30, 35, 40, 44, 50, 65,

66, 67, 68, 69, 71, 156, 203, Pl. 86 ssp. teres 157 var. tenellum 157 var. teres 157squarrulosum 157steerei 37, 47, 67, 75, 76, 96, 343, Pl. 5stewartii 85stollei 344strictum 29, 33, 47, 56, 66, 68, 69, 70, 91, 130, 133,

Pl. 39ssp. pappeanum 90stuhlmannii 339subaciphyllum 149subacutifolium 39, 67, 151subaequifolium 238subbalticum 64, 70, 340, 373, Pl. 195subbicolor 80, 83subbrachycladum 87subcontortum 237subcuspidatum 334, 335subditivum 66, 71, 336, 351, 393, Pl. 215subdivisum 145, 146suberythrocalyx 80, 87subfalcatulum 63, 71, 336, 341, 373, Pl. 195

subfulvum 22, 34, 39, 48, 66, 69, 137, 138, 140, 173, Pl. 56

ssp. purpureum 140subhomophyllum 60, 70, 230, 298, Pl. 143submedium 55, 70, 88, 125, Pl. 34submolliculum 237, 336submollissimum 244submolluscum 244submucronatum 211subnitens 12, 15, 16, 22, 35, 40, 44, 50, 58, 65, 66, 67,

68, 69, 70, 71, 138, 140, 144, 151, 153, 154, 197, Pl. 80

var. fl avicomans 154 var. nitidum 140 var. obscurum 154 var. patulum 139subobesum 41, 67, 240, 241, 320, Pl. 165suborbiculare 224subovalifolium 60, 70, 241, 318, Pl. 163subrecurvum 344 var. borneense 344subrigidum 144subrotundifolium 207subrufescens 60, 70, 241, 318, Pl. 163subsecundoides 15, 56, 70, 81, 82, 111, Pl. 20subsecundum 12, 13, 14, 21, 22, 30, 31, 35, 41, 45, 61,

65, 66, 67, 68, 69, 70, 147, 216, 225, 226, 290, Pl. 135

var. andrusii 212 var. carolinianum 234 var. contortum 234 var. inundatum 218 var. isophyllum 235 var. latissimum 238 var. platyphyllum 238subserratum 63, 71, 336, 341, 373, Pl. 195subtile 16, 144subtursum 80subulatum 132, 144subundulatum 344sucrei 59, 70, 233, 305, Pl. 150sulcatum 217, 220sullivanii 237sullivantianum 75sullivantii 75sulphureum 80sumapazense 59, 70, 243, 327, Pl. 172surisici 338tabulare 135 var. molluscoides 135tabuleirense 15, 87takedae 236talbotianum 138, 139tenellum 13, 14, 15, 22, 27, 31, 37, 43, 46, 54, 65, 66,

67, 68, 69, 70, 71, 138, 157, 352, 396, Pl. 218tenerum 34, 49, 66, 69, 135, 136, 140, 141, 144, Pl. 58tenue 225tenuifolium 137

eschweizerbart_XXX

Sample

page

s

Index 435

teres 12, 13, 14, 15, 21, 22, 35, 40, 50, 66, 67, 69, 157, 204, Pl. 87

var. concinnum 132 var. squarrosum 156tescorum 38, 48, 67, 69, 134, 135, 163, Pl. 46thailandense 239thomsonii 152tijucae 85, 87tonduzii 146torreyanum 53, 69, 351, 395, Pl. 217torreyi 351tosaense 42, 67, 68, 351, 352, 388, Pl. 210trachyacron 80trachynotum 83transvaaliense 226trichophyllum 335tricladum 147trigonum 244trinitense 53, 64, 69, 70, 71, 341, 344, 345, 376, Pl. 198 var. fi tzgeraldii 336triporosum 137trirameum 51, 69, 226, 298, Pl. 134tristichum 90troendelagicum 36, 66, 342, 377, Pl. 199trollii 15, 55, 69, 89, 126, Pl. 35truncatum 46, 68, 226, 227, 236, Pl. 136, 137, 291, 292 var. bordasii 226, 227, 292, Pl. 117 var. truncatum 226, 227, 291, Pl. 116tumidulum 31, 68, 211, 258, 259, Pl. 103, 104 var. confusum 44, 259 var. macrophyllum 211 var. microphyllum 211 var. tumidulum 44, 258tundrae 35, 40, 50, 66, 67, 69, 157, 205, Pl. 88turfaceum 74turgens 63, 70, 227, 288, Pl. 133turgescens 60, 70, 243, 318, Pl. 163turgidulum 215turgidum 215tursum 85typicum 66ugandense 208, 248uleanum 62, 70, 227, 293, Pl. 138umbrosum 21, 61, 70, 227, 294, Pl. 139undulatum 335uruguayense 62, 70, 228, 294, Pl. 139usteri 135 var. versicolor 153 var. viride 142uzenense 240, 241validum 218, 220vancouverense 137vandenbroeckii 208

variabile 344 var. intermedium 338, 347 var. majus 347variegatum 225venustum 34, 48, 66, 69, 141, 174, Pl. 57veresczagini 133versicolor 142versiporum 59, 70, 241, 320, Pl. 142vesiculare 85violascens 39, 44, 67, 68, 141, 175, 176, Pl. 58, 59virginianum 344viride 344, 345vitalii 55, 70, 89, 126, Pl. 35vitianum 87vitjianum 80vogesiacum 338vulcanicum 90vulgare 80waghornei 78wallisii 85wardellense 83warnstorfi anum 142warnstorfi i 15, 21, 22, 34, 39, 49, 66, 67, 69, 133, 139,

142, 151, 177, Pl. 60 var. pallens 150 var. pseudopatulum 150wattsii 335weberbaueri 149weberi 344weddelianum 85wenckei 207weymouthii 89wheeleri 90whiteleggei 83wieboldtii 52, 69, 342, 394, Pl. 216wilcoxii 83wilfi i 49, 69, 151, 191, Pl. 74wilsonii 138, 142 var. roseum 139wrightii 87wulfi anum 7, 14, 15, 20, 21, 22, 27, 30, 35, 40, 51, 66,

67, 69, 157, 206, Pl. 89wulfi i 157xerophilum 218, 220zickendrathii 338

Takakia lepidozioides 20Usnea 11Vorcutannularia laevis 17 plicata 17

eschweizerbart_XXX

Sphagnum specialist Dierk Michaelis documents the worldwide known peat moss species (genus Sphagnum) and presents keys for their identification. It represents the updated, supplemented English language version of the author’s original peat moss flora of 2011 (in German), the first overall presenta-tion of Sphagnum since Carl Warnstorf’s “Sphag-nologia Universalis” of 1911. Compared to the Ger-man edition, 12 species have been added, 23 new plates were added, the chapters on phylogeny and research history have been revised and a new chap-ter on Sphagnum ecology has been added.Since Warnstorf’s comprehensive work, numerous names have been recognized and revised as syn-onyms - particularly by Andrews, Eddy and Isoviita. These revisions, and the approximately 150 new species described since then, have been incorpo-rated into this volume, as well as the results of the author’s own studies. Genetic characteristics were used to define the species of problematic groups.The peat mosses are of key ecological and econom-ic importance among the mosses. They populate al-most all continents with a clear focus on northern South America, North America, East and North Asia and Europe.The genus Sphagnum is very isolated within the Bryopsida, similarities in the construction of the sporophyte indicate a distant relation to the rock-mosses (class Andreaeopsida). For the internal classification of Sphagnum there are very different approaches with up to 4 subgen-era and up to 18 possible sections, of which 14 are distinguished in this volume.

Peat mosses in the narrow sense (genus Sphag-num) feature a combination of leaf dimorphism (stem and branch leaves), cell dimorphism (living chlorophyll and empty hyaline cells) and branch di-morphism (strongly assimilating spreading branch-es and hanging branches serving the outer water supply) that is unique among mosses. Although the assignment of any peat moss to the genus Sphag-num usually does not cause any problems, the de-termination down to the species level causes dif-ficulties sometimes.The author introduces and describes the anatomy and morphology of Sphagnum, and explains the reproductive biology, the research history and phy-logenesis of peat mosses. The systematic part is divided into three segments: Description and identification of the sections, keys for all peat moss species, separated by continents, as well as Sphagnum species lists for 20 phytogeographic regions of the world.The keys for Africa, Europe and North America are based on existing data and were revised and supple-mented with the help of recent descriptions, updated species concepts and new floristic data. Completely new keys have been developed for South America and Asia, as these did not exist previously. 292 peat moss species are described in detail, supplemented by data on habitats, geographical distribution and lists of synonyms.This section is supplemented by the presentation of the inner and outer characteristics on 219 plates. A very extensive bibliography rounds off the volume.

www.schweizerbart.deISSN 0067-7892

D. Michaelis, The Sphagnum Species of the WorldBibliotheca Botanica, Volume 162, 2019

9 783510 480333

ISBN 978-3-510-48033-3

eschweizerbart_XXX

Sphagnum

Johannesstr. 3A, 70176 Stuttgart, Germany., Tel. +49 (0)711 351456-0, Fax +49 (0)711 351456-99, order@schweizerbart.de, www.schweizerbart.de

Original Contributions to BotanyOriginal Contributions to Botany, founded in 1886 www.schweizerbart.de/series/biblbotanica ISSN 0067-7892

Since Warnstorf’s comprehensive work, numerous names have been recognized and revised as syn-onyms - particularly by Andrews, Eddy and Isoviita. These revi-sions, and the approximately 150 new species described since then, have been incorporated into this volume, as well as the results of the author’s own studies. Genetic characteristics were used to define the species of problematic groups.The peat mosses are of key eco-logical and economic importance among the mosses. They populate almost all continents with a clear focus on northern South America, North America, East and North Asia and Europe.The genus Sphagnum is very iso-lated within the Bryopsida, similari-ties in the construction of the spo-rophyte indicate a distant relation to the rock-mosses (class Andre-aeopsida). For the internal classification of Sphagnum there are very different

approaches with up to 4 subgen-era and up to 18 possible sections, of which 14 are distinguished in this volume.Peat mosses in the narrow sense (genus Sphagnum) feature a combination of leaf dimorphism (stem and branch leaves), cell di-morphism (living chlorophyll and empty hyaline cells) and branch dimorphism (strongly assimilating spreading branches and hanging branches serving the outer wa-ter supply) that is unique among mosses. Although the assignment of any peat moss to the genus Sphagnum usually does not cause any problems, the determination down to the species level causes difficulties sometimes.The author introduces and de-scribes the anatomy and morphol-ogy of Sphagnum, and explains the reproductive biology, the re-search history and phylogenesis of peat mosses. The systematic part is divided into three segments:

Description and identification of the sections, keys for all peat moss species by continent, and Sphagnum species lists for 20 phytogeographic regions of the world.

The keys for Africa, Europe and North America are based on ex-isting data and were revised and supplemented using recent de-scriptions, updated species con-cepts and new floristic data. Com-pletely new keys were developed for South America and Asia, as these did not exist previously. 292 peat moss species are described in detail, supplemented by data on habitats, geographical distribution and lists of synonyms.This section is supplemented by the presentation of the inner and outer characteristics on 219 plates. A very extensive bibliography rounds off the volume.

Sphagnum specialist Dierk Michaelis documents the worldwide known peat moss species (genus Sphagnum) and presents keys for their identifi-cation. It represents the updated, augmented English language version of the author’s original peat moss flora of 2011 (in German), the first overall presentation of Sphagnum since Carl Warnstorf’s “Sphagnologia Univer-salis” of 1911. Compared to the German edition, 12 species have been added, as were 23 new plates; the chapters on phylogeny and research history were revised and a new chapter on Sphagnum ecology was added.

Sphagnum

Sphagnum

Bibliotheca Botanica 162. The Bibliotheca Botanica 162. The SphagnumSphagnum Species of the World Species of the World

Sphagnum Acocosphagnum. . . . . phagnum Inretorta . . . . . . . . . . .

Sphagnum SphagnumSphagnum Rigida . . . . . . . . . . . . . Sphagnum Insulosa . . . . . . . . . . . Sphagnum Acutifolia . . . . . . . . . . Sphagnum Squarrosa. . . . . . . . . Sphagnum Polyclada . . . . . . . . . Sphagnum Acrosphagnum . . . . .

Sphagnum Subsecunda . . . . . . Sphagnum Isocladus. . . . . . . . . Sphagnum Hemitheca . . . . . . . Sphagnum Cuspidata . . . . . . . . Sphagnum Mollusca . . . . . . . . . Ambuchanania. . . . . . . . . . . . . . . . . .

143. Gabriele Kothe-Heinrich: Revision der Gattung Ha-lothamnus (Chenopodiaceae). Dissertation Kassel 1992. 1993. 176 S., 73 Abb., 12 Tab., 2 Photos, 12 Phototaf., ISBN 978-3-510-48014-2, brosch.

144. Dagmar Lange: Untersuchungen zur Systematik und Taxonomie der Gattung Helictotrichon Besser ex J.A. Schultes & J. H. Schultes (Poaceae) in Südosteuropa und Vorderasien. 1995. IV, 238 S., 49 Abb., 16 Tab., ISBN 978-3-510-48015-9, brosch.

145. Christoph Weiglin: Freilandökologische Untersu-chungen und Gewächshausexperimente zur Ef zienz hygroskopisch beweglicher Diasporen von Sperma-tophyten. 1995. VIII, 160 S., 62 Abb., 44 Tab., ISBN 978-3-510-48016-6, brosch.

146. R. Ch. Kruijt: A taxonomic monograph of Sapium Jacq., Anomostachys (Baill.) Hurus., Duvigneaudia J. Léonard and Sclerocroton Hochst. (Euphorbiaceae tribe Hippomaneae). 1996. V, 109 p., 35 gs., 8 tab., ISBN 978-3-510-48017-3, paperback.

147. Ewald Gerhardt: Taxonomische Revision der Gat-tungen Panaeolus and Panaeolina (Fungi, Agarica-les, Coprinaceae). 1996. 149 S., 77 Abb., ISBN 978-3-510-48018-0, brosch.

148. Jens G. Rohwer: Die Frucht- und Samenstrukturen der Oleaceae. Eine vergleichend-anatomische Unter-suchung. 1996. V, 177 S., 300 Abb., 6 Tab., ISBN 978-3-510-48019-7, brosch.

149. Sabrina Rilke: Revision der Sektion Salsola S.L. der Gattung Salsola (Chenopodiaceae). 1999. 189 S., 75 Abb., 7 Tab., ISBN 978-3-510-48020-3, brosch.

150. Ragna Mißkampf und Wiebke Züghart: Floristisch-ökologische Untersuchung der Spontan ora in Bre-mer Häfen unter besonderer Berücksichtigung der anthropochoren P anzen. 2000. 110 S., 25 Abb., 38 Tab., 10 Photos, ISBN 978-3-510-48021-0, brosch.

151. Joachim Prutsch und Rainer Schill: Die Ontogenese der Narbe bei den Orchideen. 2000. 82 S., 2 Abb., 1 Tab., 249 Taf., ISBN 978-3-510-48022-7, brosch.

152. Iris Mundry: Morphologische und morphogenetische Untersuchungen zur Evolution der Gymnospermen. 2000. 90 S., 72 Abb., ISBN 978-3-510-48023-4, brosch.

153. Christine Ehrhart: Die Gattung Calceolaria (Scrophu-lariaceae) in Chile. 2000. 283 S., 145 Abb., 17 Tab., ISBN 978-3-510-48024-1, brosch.

154. Markus Sonnberger: Aspekte der Reproduktions-biologie des Echten Steinsamen (Lithospermum of -cinale, Boraginaceae). 2002. XIII, 150 S., 13 Abb., 43 Tab., 27 Diagr., ISBN 978-3-510-48025-8, brosch.

155. Martin Schmid: Morphologie, Vergesellschaftung, Ökologie, Verbreitung und Gefährdung der Sumpf-Löwenzähne (Taraxacum sect. Palustria Dahlst., Asteraceae) Süddeutschlands. 2003. IX, 268 S., 86 Abb., 14 Tab., 25 Kart., ISBN 978-3-510-48026-5, brosch.

156. Peter Sack: Ausbreitungsbiologische Experimente an Arten der Subtribus Prunellineae (Prunella L. und Cleonia L.; Lamiaceae). 2003. XI, 121 S., 49 Abb., 25 Tab., 18 Taf., ISBN 978-3-510-48027-2, brosch.

157. Lieselotte Klingenberg: Monographie der südameri-kanischen Gattungen Haplopappus Cass. und Noto-pappus L. Klingenberg (Asteraceae – Astereae). 2007. X, 331 S., 230 Abb., 30 Tab., ISBN 978-3-510-48028-9, brosch.

158. Monika Langlotz: Aspekte der Reproduktionsbiologie chorikarper Blütenp anzen. 2007. X, 276 S., 59 Abb., 26 Graf., zahlr. Ill., ISBN 978-3-510-48029-6, brosch.

159. Arno Wörz: Revision of Eryngium L. (Apiaceae-Sa-niculoideae): General part and Palaearctic species. 2011. 498 pages, 84 gures, 41 tables, 12 plates, ISBN 978-3-510-48030-2, bound

160. Dierk Michaelis: Die Sphagnum-Arten der Welt. 2011. 408 Seiten, 10 Abbildungen, 194 Tafeln. ISBN 978-3-510-48031-9, gebunden

161. Veit Martin Dörken: The evolutionary relevance of vegetative long-shoot/short-shoot differentiation in gymnospermous tree species. 2012. 100 pages, 41 gures, 12 tables. ISBN 978-3-510-48032-6, paperback