Atlas of Wood, Bark and Pith Anatomy of Eastern Mediterranean Trees and Shrubs

Atlas of Wood, Bark and Pith Anatomy of Eastern Mediterranean Trees and Shrubswith a Special Focus on Cyprus

Dr. Alan CrivellaroUniversità degli Studi di Padova – Dipartimento Territorio e Sistemi AgroForestaliViale dell’Università 1635020 Legnaro (PD), Italyemail: alancrivellaro@gmail.com

Prof. Dr. Fritz Hans SchweingruberInstitute for Forest, Snow and Landscape Research – WSLZürcherstrasse 1118903 Birmensdorf, Switzerlandemail: fritz.schweingruber@wsl.ch

Charalambos S. ChristodoulouTakis PapachristophorouTakis TsintidesDept. of Forests, Ministry of Agriculture, Natural Resources and EnvironmentLouki Akrita 261414 Nicosia, Cyprus

Alessia Da RosGraphic Designer

ISBN 978-3-642-37234-6 ISBN 978-3-642-37235-3 (eBook)DOI 10.1007/978-3-642-37235-3Springer Heidelberg New York Dordrecht London Library of Congress Control Number 2013942798 © Springer-Verlag Berlin Heidelberg 2013This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recita-tion, broadcasting, reproduction on microfilms or in any other physical way, and transmission or infor-mation storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar meth-odology now known or hereafter developed. Exempted from this legal reservation are brief excerpts in connection with reviews or scholarly analysis or material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Duplica-tion of this publication or parts thereof is permitted only under the provisions of the Copyright Law of the Publisher’s location, in its current version, and permission for use must always be obtained from Springer. Permissions for use may be obtained through RightsLink at the Copyright Clearance Center. Violations are liable to prosecution under the respective Copyright Law. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and reg-ulations and therefore free for general use. While the advice and information in this book are believed to be true and accurate at the date of publication, neither the authors nor the editors nor the publisher can accept any legal responsibility for any errors or omissions that may be made. The publisher makes no warranty, express or implied, with respect to the material contained herein.

Cover illustrationCross-section of a twig of Phlomis lunariifolia (Lamiaceae), grown in the maquis on Cyprus. Cells of the central part of the pith are thin-walled and tracheary elements of the metaxylem are arranged in distinct radial rows. Printed on acid-free paper

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Alan Crivellaro received his BA in Wood Science, a MS in Forestry and a PhD in Ecology from the University of Pa-dova, Italy. Timber strength properties and wood anatomy are a particular focus of his work. For this book he collect-ed the material, prepared the slides, realized the anatomi-cal descriptions and photomicrographs, and supervised the graphical design of the atlas.

Fritz H. Schweingruber has a MS in Botany (Bern Universi-ty, Switzerland). He was leader of the research group Tree Ring and Site at the Swiss Federal Institute for Forests, Snow and Landscape Research (WSL) and professor at the Institute of Botany at the University of Basel, Switzerland. His work is documented by many scientific papers and wood anatomy books. He supervised the material prepara-tion, anatomical descriptions, and atlas’ design.

Takis Tsintides has a BS in Forestry (Aristotle University, Greece) and an MS in Recourse Management (Edinburgh University, Scotland). He organised and supervised the field-work for the collection of samples on Cyprus and coordinated the editing of plant descriptions.

Charalambos S. Christodoulou has a BS in Forest Man-agement (University of Central Lancashire, UK). He col-lected and identified plant material, provided plant photographs and contributed to the editing of plant de-scriptions.

Takis Papachristophorou has a Diploma in Forestry from the Cyprus Forestry College and a Diploma in Tree Surgery (Germany). He collected and identified plant material and provided photographs of plants.

Alessia Da Ros has a BA in Industrial Design and a MS in Visual and Multimedia Communications at the University of Venice, Italy. She worked in a graphic studio; at the moment she is in charge of organizing art exhibitions. For this book she managed the graphic design, along with text and photos editing.

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The idea of this atlas started in 2004 after Alan Crivellaro visited the island of Cyprus and bought a book about tree and shrub spe-cies growing on the island. Soon after return-ing home, he realized that the wood anato-mies were only described for a portion of the species included in that book. Several months later Alan Crivellaro and Fritz Schweingru-ber became acquainted at the International Course on Wood Anatomy and Tree-Ring Ecol-ogy in Davos Laret (Switzerland). They dis-cussed the feasibility of a large study focused on the anatomy of Cyprus’ trees and shrubs, with the recognition that the indigenous flora of the island can be considered representative of the eastern Mediterranean region. It wasn’t until 2009 that they started collecting plant material, with the essential assistance of their Cypriot coauthors, Charalambos S. Christo-doulou, Takis Papachristophorou and Takis Tsintides. The collecting was followed by slides preparation, material description, and finally preparation of this atlas.This book is intended to serve as a manual for the identification of stem material under the microscope for all woody plants (trees, shrubs, subshrubs and climbers), whether wild or cultivated, that are commonly found in the Eastern Mediterranean region. Along with the anatomical descriptions of stem wood, the atlas describes also the anatomical structure of twigs’ xylem, bark and pith. These addi-tional information will aid in plant identifica-tion, and also will open the way for new types of dendroecological and functional studies.

Preface and Acknowledgments

We hereby acknowledge the following people and organizations for their contributions to this book: Tiziana Urso (Padova, Italy) who introduced AC to wood anatomy; Maria Socra-tous and Andreas Nearchou (Nicosia, Cyprus) who helped during some field collection trips; Holger Gärtner and Paolo Cherubini (Birmens-dorf, Switzerland) for hosting AC at WSL and making substantial contributions during the formative stages of the project; Petra Zi-bulski (Basel, Switzerland) who made many important critical remarks and suggestions to improve codified lists for bark and pith ana-tomical features; Barbara Lachenbruch (Cor-vallis, Oregon) for hosting AC at OSU and for the stimulating discussions providing critical scientific advices and valuable help in the preparation of the manuscript; Annett Börner (Adelaide, Australia), Andrew Brookes (Ports-mouth, United Kingdom), Shaaban Ghaland-arayeshi (Kordkoy, Iran), Ori Fragman-Sapir (Jerusalem, Israel) and Georgios N. Hadjik-yriacou (Nicosia, Cyprus) who provided plant pictures; and Alma Piermattei (Ancona, Italy) who provided some specimens and plant pictures.

The Cyprus’ Ministry of Agriculture, Natural Resources and Environment - Department of Forests, the University of Padova (Italy) - De-partment of Land, Environment, Agriculture, and Forestry (TeSAF), and the Swiss Federal Institute for Forest, Snow and Landscape Re-search (WSL, Birmensdorf, Switzerland) pro-vided logistic and scientific infrastructures.

Alan Crivellaro and Fritz H. SchweingruberPadova and Birmensdorf, January 2013

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Humans have probably always lived with some balance between admiration of the beauty in nature and reliance on what nature can provide us. Wood is a good example. Woodworkers, carvers, builders and paper-makers have created objects and shelters that celebrate wood, exuberantly or quietly. Simultaneously, half the harvested wood on earth is used for fuel to make dwellings habitable and food edible, and most of us use wood products for daily needs from furni-ture to paper. It is estimated that the aver-age person on earth uses the equivalent of 1,6 kg of dried wood a day—about 600 kg in a year—with the highest consumers, in the US, using four times that much. Economists argue that if we are to continue raising the standard of living throughout the world we need to use more wood, not less, because its use is less costly overall than the alternative materials such as aluminium and steel. Most people consider the stem as “just a piece of wood” when in fact, being situated mid-way between roots and canopy, its activities are essential for the functioning of the plant. The stem transports water, nutrients, sugars, and hormones; buffers water uptake and loss; supports the mass of the canopy plus loads from wind, snow, ice, fruit, and epiphytes; displays foliage and flowers to resources like light and pollinators; protects the plant from abrasion and fire; and harbors materials and meristems that may be needed for growth, reproduction, and recovery from disturbance. All this is done by an organ that is fully func-tional at the same time that it is able to grow. An enormous number of ways have evolved to perform these functions, and in consequence, there is an enormous diversity of types of woods and barks. This Atlas of Wood, Bark and Pith Anatomy of Eastern Mediterranean Trees and Shrubs is a beautiful contribution toward understanding stems and their adap-tations. Moreover, the displays of excellently prepared and stained sections are art in their own right. This atlas provides a rich treasury of infor-mation on stems. Unlike most compendia, this book covers an entire woody flora at one location, and thus gives us a glimpse of the variety of stem tissues that can co-occur.

Foreword

The island of Cyprus is sufficiently diverse geographically and botanically that we can use this atlas to look for patterns in the tis-sues related to phylogeny, growth form, and level of endemism. Another unusual feature of this book is the inclusion of images and descriptions of three plant parts in addition to the main stem: twig xylem, the pith region, and the twig’s bark region. The twig/stem juxtapositions show us graphically that in many cases twigs are quite distinct anatomi-cally and are not just miniature trunks. The pith images and descriptions detail the sur-prising diversity of pith anatomies in terms of overall morphology, the variety of cell types within one pith, the shape of the pith cells, and the thickness and optical properties of their walls. The region just exterior to the pith illustrates the primary xylem and the earliest-developed secondary xylem, provid-ing a snapshot of structures present during the vulnerable development stage just after stem elongation. The images of bark dem-onstrate the many ways it is constructed in twigs. Lastly, having the bark, twig, pith, and stem in a tidy two-page spread invites us to make comparisons among them. We can look, for example, at whether thick-walled fibers, thick-walled pith parenchyma, and sclereids are all found in the same plant or whether these wall thicknesses are uncoupled from tissue to tissue and organ to organ. The user will also appreciate the plant descriptions and photographs, which help us link the micro-scopic views to the scale at which we may have known the plants in our hikes and in some cases, in our gardens and kitchens. An understanding of the stem’s adaptations is of growing importance for management. Humans are increasingly relying on plants, but in the future these plants are likely to be growing in more marginal habitats and in cli-mates with more frequent droughts, flooding, and wind. Humans are also finding new uses for species that we have used for millennia, such as phytoremediation, bioenergy, carbon sequestration, and erosion control. Manager and breeders need guidance on what traits—the structures and functions together—con-tribute to success in the type of habitat where they will be grown. Unfortunately, many of

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the biologists, agronomists, and foresters involved in tree breeding and selection are no longer trained in anatomy, and instead of in-vestigating the structure/function complexes, they tend to use proxies (like wood density) to estimate the plant’s success.This atlas can fill part of this gap by serving to educate us in the many structural variants that survive and reproduce in one geographic area. For some of the atlas’ users, this case study of anatomical diversity will provide a database to help them study how plants work.

Such research is helping increase the number of scientifically based tales on our botani-cal bookshelves and decrease the number of “Just So” stories (named for Rudyard Kipling’s children’s stories in which the narrator tells an unverified purpose for a biological trait). Other readers will use this atlas for specific information on the species or on in the cli-mates encompassed here. Still other readers will enjoy the book for its beauty. However it is used, this atlas is a welcome addition to the resources we have on plant anatomy.

Barbara LachenbruchDept. of Forest Ecosystems and Society,

Oregon State UniversityCorvallis, Oregon, USA

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Preface and Acknowledgments..............VIIForeword..............................................IX

1. Introduction..........................................12. Materials and Methods............................3

Origin of the material studied..................3Plant material preparation.......................3Wood density..........................................3

3. Definition of Anatomical Features............5Xylem..................................................5Bark..................................................24Pith...................................................31

4. Identification Keys.................................37Wood without vessels (Gymnosperms)...38Wood with vessels and included phloem...39Wood with vessels and without includedphloem..................................................40

5. Anatomical Description of Species........49Gymnosperms................................53 Cupressaceae............................54 Pinaceae....................................64 Taxaceae.................................76Gnetales.......................................79 Ephedraceae..............................80Angiosperms: Monocotyledons..............85 Asparagaceae................................86 Graminaceae..............................88 Ruscaceae.....................................90 Smilacaceae..................................91Angiosperms: Dicotyledons....................93 Adoxaceae......................................94 Amaranthaceae.............................100 Anacardiaceae..............................102 Apocynaceae................................116 Aquifoliaceae................................120 Araliaceae.....................................122 Aristolochiaceae...........................124 Asclepiadaceae.............................126 Asteraceae....................................128 Berberidaceae...............................154 Betulaceae....................................156 Boraginaceae................................162 Brassicaceae.................................172 Buxaceae......................................188 Caesalpiniaceae............................190 Cannabaceae..............................192 Capparaceae...............................196 Caprifoliaceae............................198

Caryophyllaceae.........................198 Chenopodiaceae........................202 Cistaceae...................................208 Convolvulaceae..........................228 Cornaceae...................................256 Dipsacaceae..............................258 Ebenaceae..................................264 Elaeagnaceae...........................266 Ericaceae...................................268 Euphorbiaceae...........................278 Fabaceae.....................................284 Fagaceae....................................312 Frankeniaceae.............................326 Hypericaceae...............................328 Juglandaceae...............................332 Lamiaceae...................................334 Lauraceae...................................400 Lythraceae..................................402 Malvaceae..................................404 Mimosaceae..............................406 Moraceae.....................................410 Myrtaceae....................................418 Oleaceae......................................430 Orobanchaceae...........................440 Phytolaccaceae...........................442 Platanaceae................................444 Plumbaginaceae........................446 Polygonaceae...............................450 Ranunculaceae.............................452 Rhamnaceae................................456 Rosaceae.....................................466 Rubiaceae....................................506 Rutaceae......................................514 Salicaceae....................................522 Sapindaceae.................................528 Scrophulariaceae..........................536 Simarubaceae..............................538 Solanaceae...................................540 Styracaceae..................................548 Tamaricaceae...............................550 Thymelaeaceae............................558 Ulmaceae.....................................562 Verbenaceae............................564 Vitaceae........................................568 Zygophyllaceae.............................570

References.......................................575List of Species and Families................579

Table of Contents

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Map of the Mediterranean Basin

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Legend:AE East Aegean IslandsAg AlgeriaAl AlbaniaAn Asiatic TurkeyBl Balearic IslandsBu Bulgaria

Co CorsicaCr Crete and KarpathosCy CyprusEg EgyptGa FranceGr GreeceHs Spain

IJ Israel and Jordan It ItalyJu ex JugoslaviaLi LibyaLS Lebanon and SyriaLu PortugalMa Morocco

Me MaltaRK CrimeaSa SardiniaSi SicilySn Sinai Tn TunisiaTu Turkey in Europe

Range maps of the Mediterranean basin for all species. Topography is shown with con-tours at 1000 and 2000 m above sea level. Black dots indicate the country in which the species is found.

Range maps of Cyprus for all endemic species. Topography is shown with contours at 100 m above sea level. These maps also show the eight botanical divisions (sensu Meikle 1979). Red square indications Troodos Peak (eleva-tion 1952 m).

Range distribution maps