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

589 Pages·2013·270.3 MB·English

by  Alan Crivellaro

Alan Crivellaro
Fritz Hans Schweingruber
Atlas of Wood, Bark
and Pith Anatomy of
Eastern Mediterranean
Trees and Shrubs
with a Special Focus on Cyprus
Atlas of
Wood, Bark and Pith Anatomy
of Eastern Mediterranean
Trees and Shrubs
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Atlas of
Wood, Bark and Pith Anatomy
of Eastern Mediterranean
Trees and Shrubs
with a Special Focus on Cyprus
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Dr. Alan Crivellaro
Università degli Studi di Padova – Dipartimento Territorio e Sistemi AgroForestali
Viale dell’Università 16
35020 Legnaro (PD), Italy
email: [email protected]
Prof. Dr. Fritz Hans Schweingruber
Institute for Forest, Snow and Landscape Research – WSL
Zürcherstrasse 111
8903 Birmensdorf, Switzerland
email: [email protected]
Charalambos S. Christodoulou
Takis Papachristophorou
Takis Tsintides
Dept. of Forests, Ministry of Agriculture, Natural Resources and Environment
Louki Akrita 26
1414 Nicosia, Cyprus
Alessia Da Ros
Graphic Designer
ISBN 978-3-642-37234-6 ISBN 978-3-642-37235-3 (eBook)
DOI 10.1007/978-3-642-37235-3
Springer Heidelberg New York Dordrecht London

Library of Congress Control Number 2013942798

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Cover illustration
Cross-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
Springer is part of Springer Science+Business Media (www.springer.com)
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.
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.
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.
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.
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.
V
Preface and Acknowledgments
The idea of this atlas started in 2004 after We hereby acknowledge the following people
Alan Crivellaro visited the island of Cyprus and organizations for their contributions to
and bought a book about tree and shrub spe- this book: Tiziana Urso (Padova, Italy) who
cies growing on the island. Soon after return- introduced AC to wood anatomy; Maria Socra-
ing home, he realized that the wood anato- tous and Andreas Nearchou (Nicosia, Cyprus)
mies were only described for a portion of the who helped during some field collection trips;
species included in that book. Several months Holger Gärtner and Paolo Cherubini (Birmens-
later Alan Crivellaro and Fritz Schweingru- dorf, Switzerland) for hosting AC at WSL and
ber became acquainted at the International making substantial contributions during the
Course on Wood Anatomy and Tree-Ring Ecol- formative stages of the project; Petra Zi-
ogy in Davos Laret (Switzerland). They dis- bulski (Basel, Switzerland) who made many
cussed the feasibility of a large study focused important critical remarks and suggestions to
on the anatomy of Cyprus’ trees and shrubs, improve codified lists for bark and pith ana-
with the recognition that the indigenous flora tomical features; Barbara Lachenbruch (Cor-
of the island can be considered representative vallis, Oregon) for hosting AC at OSU and for
of the eastern Mediterranean region. It wasn’t the stimulating discussions providing critical
until 2009 that they started collecting plant scientific advices and valuable help in the
material, with the essential assistance of their preparation of the manuscript; Annett Börner
Cypriot coauthors, Charalambos S. Christo- (Adelaide, Australia), Andrew Brookes (Ports-
doulou, Takis Papachristophorou and Takis mouth, United Kingdom), Shaaban Ghaland-
Tsintides. The collecting was followed by arayeshi (Kordkoy, Iran), Ori Fragman-Sapir
slides preparation, material description, and (Jerusalem, Israel) and Georgios N. Hadjik-
finally preparation of this atlas. yriacou (Nicosia, Cyprus) who provided plant
This book is intended to serve as a manual pictures; and Alma Piermattei (Ancona, Italy)
for the identification of stem material under who provided some specimens and plant
the microscope for all woody plants (trees, pictures.
shrubs, subshrubs and climbers), whether
wild or cultivated, that are commonly found in The Cyprus’ Ministry of Agriculture, Natural
the Eastern Mediterranean region. Along with Resources and Environment - Department of
the anatomical descriptions of stem wood, the Forests, the University of Padova (Italy) - De-
atlas describes also the anatomical structure partment of Land, Environment, Agriculture,
of twigs’ xylem, bark and pith. These addi- and Forestry (TeSAF), and the Swiss Federal
tional information will aid in plant identifica- Institute for Forest, Snow and Landscape Re-
tion, and also will open the way for new types search (WSL, Birmensdorf, Switzerland) pro-
of dendroecological and functional studies. vided logistic and scientific infrastructures.
Alan Crivellaro and Fritz H. Schweingruber
Padova and Birmensdorf, January 2013
VII
Foreword
Humans have probably always lived with The island of Cyprus is sufficiently diverse
some balance between admiration of the geographically and botanically that we can
beauty in nature and reliance on what nature use this atlas to look for patterns in the tis-
can provide us. Wood is a good example. sues related to phylogeny, growth form, and
Woodworkers, carvers, builders and paper- level of endemism. Another unusual feature
makers have created objects and shelters of this book is the inclusion of images and
that celebrate wood, exuberantly or quietly. descriptions of three plant parts in addition
Simultaneously, half the harvested wood to the main stem: twig xylem, the pith region,
on earth is used for fuel to make dwellings and the twig’s bark region. The twig/stem
habitable and food edible, and most of us use juxtapositions show us graphically that in
wood products for daily needs from furni- many cases twigs are quite distinct anatomi-
ture to paper. It is estimated that the aver- cally and are not just miniature trunks. The
age person on earth uses the equivalent of pith images and descriptions detail the sur-
1,6 kg of dried wood a day—about 600 kg in prising diversity of pith anatomies in terms of
a year—with the highest consumers, in the overall morphology, the variety of cell types
US, using four times that much. Economists within one pith, the shape of the pith cells,
argue that if we are to continue raising the and the thickness and optical properties of
standard of living throughout the world we their walls. The region just exterior to the
need to use more wood, not less, because its pith illustrates the primary xylem and the
use is less costly overall than the alternative earliest-developed secondary xylem, provid-
materials such as aluminium and steel. Most ing a snapshot of structures present during
people consider the stem as “just a piece of the vulnerable development stage just after
wood” when in fact, being situated mid-way stem elongation. The images of bark dem-
between roots and canopy, its activities are onstrate the many ways it is constructed in
essential for the functioning of the plant. The twigs. Lastly, having the bark, twig, pith, and
stem transports water, nutrients, sugars, and stem in a tidy two-page spread invites us to
hormones; buffers water uptake and loss; make comparisons among them. We can look,
supports the mass of the canopy plus loads for example, at whether thick-walled fibers,
from wind, snow, ice, fruit, and epiphytes; thick-walled pith parenchyma, and sclereids
displays foliage and flowers to resources like are all found in the same plant or whether
light and pollinators; protects the plant from these wall thicknesses are uncoupled from
abrasion and fire; and harbors materials and tissue to tissue and organ to organ. The user
meristems that may be needed for growth, will also appreciate the plant descriptions and
reproduction, and recovery from disturbance. photographs, which help us link the micro-
All this is done by an organ that is fully func- scopic views to the scale at which we may
tional at the same time that it is able to grow. have known the plants in our hikes and in
An enormous number of ways have evolved to some cases, in our gardens and kitchens.
perform these functions, and in consequence, An understanding of the stem’s adaptations
there is an enormous diversity of types of is of growing importance for management.
woods and barks. This Atlas of Wood, Bark Humans are increasingly relying on plants,
and Pith Anatomy of Eastern Mediterranean but in the future these plants are likely to be
Trees and Shrubs is a beautiful contribution growing in more marginal habitats and in cli-
toward understanding stems and their adap- mates with more frequent droughts, flooding,
tations. Moreover, the displays of excellently and wind. Humans are also finding new uses
prepared and stained sections are art in their for species that we have used for millennia,
own right. such as phytoremediation, bioenergy, carbon
This atlas provides a rich treasury of infor- sequestration, and erosion control. Manager
mation on stems. Unlike most compendia, and breeders need guidance on what traits—
this book covers an entire woody flora at one the structures and functions together—con-
location, and thus gives us a glimpse of the tribute to success in the type of habitat where
variety of stem tissues that can co-occur. they will be grown. Unfortunately, many of
IX
the biologists, agronomists, and foresters Such research is helping increase the number
involved in tree breeding and selection are no of scientifically based tales on our botani-
longer trained in anatomy, and instead of in- cal bookshelves and decrease the number of
vestigating the structure/function complexes, “Just So” stories (named for Rudyard Kipling’s
they tend to use proxies (like wood density) children’s stories in which the narrator tells
to estimate the plant’s success. an unverified purpose for a biological trait).
This atlas can fill part of this gap by serving Other readers will use this atlas for specific
to educate us in the many structural variants information on the species or on in the cli-
that survive and reproduce in one geographic mates encompassed here. Still other readers
area. For some of the atlas’ users, this case will enjoy the book for its beauty. However it
study of anatomical diversity will provide a is used, this atlas is a welcome addition to the
database to help them study how plants work. resources we have on plant anatomy.
Barbara Lachenbruch
Dept. of Forest Ecosystems and Society,
Oregon State University
Corvallis, Oregon, USA
X
Table of Contents
Preface and Acknowledgments..............VII Caryophyllaceae.........................198
Foreword..............................................IX Chenopodiaceae........................202
1. Introduction..........................................1 Cistaceae...................................208
2. Materials and Methods............................3 Convolvulaceae..........................228
Origin of the material studied..................3 Cornaceae...................................256
Plant material preparation.......................3 Dipsacaceae..............................258
Wood density..........................................3 Ebenaceae..................................264
3. Definition of Anatomical Features............5 Elaeagnaceae...........................266
Xylem..................................................5 Ericaceae...................................268
Bark..................................................24 Euphorbiaceae...........................278
Pith...................................................31 Fabaceae.....................................284
4. Identification Keys.................................37 Fagaceae....................................312
Wood without vessels (Gymnosperms)...38 Frankeniaceae.............................326
Wood with vessels and included phloem...39 Hypericaceae...............................328
Wood with vessels and without included Juglandaceae...............................332
phloem..................................................40 Lamiaceae...................................334
5. Anatomical Description of Species........49 Lauraceae...................................400
Gymnosperms................................53 Lythraceae..................................402
Cupressaceae............................54 Malvaceae..................................404
Pinaceae....................................64 Mimosaceae..............................406
Taxaceae.................................76 Moraceae.....................................410
Gnetales.......................................79 Myrtaceae....................................418
Ephedraceae..............................80 Oleaceae......................................430
Angiosperms: Monocotyledons..............85 Orobanchaceae...........................440
Asparagaceae................................86 Phytolaccaceae...........................442
Graminaceae..............................88 Platanaceae................................444
Ruscaceae.....................................90 Plumbaginaceae........................446
Smilacaceae..................................91 Polygonaceae...............................450
Angiosperms: Dicotyledons....................93 Ranunculaceae.............................452
Adoxaceae......................................94 Rhamnaceae................................456
Amaranthaceae.............................100 Rosaceae.....................................466
Anacardiaceae..............................102 Rubiaceae....................................506
Apocynaceae................................116 Rutaceae......................................514
Aquifoliaceae................................120 Salicaceae....................................522
Araliaceae.....................................122 Sapindaceae.................................528
Aristolochiaceae...........................124 Scrophulariaceae..........................536
Asclepiadaceae.............................126 Simarubaceae..............................538
Asteraceae....................................128 Solanaceae...................................540
Berberidaceae...............................154 Styracaceae..................................548
Betulaceae....................................156 Tamaricaceae...............................550
Boraginaceae................................162 Thymelaeaceae............................558
Brassicaceae.................................172 Ulmaceae.....................................562
Buxaceae......................................188 Verbenaceae............................564
Caesalpiniaceae............................190 Vitaceae........................................568
Cannabaceae..............................192 Zygophyllaceae.............................570
Capparaceae...............................196 References.......................................575
Caprifoliaceae............................198 List of Species and Families................579
XI
Map of the Mediterranean Basin
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Legend: Co Corsica IJ Israel and Jordan Me Malta
AE East Aegean Islands Cr Crete and Karpathos It Italy RK Crimea
Ag Algeria Cy Cyprus Ju ex Jugoslavia Sa Sardinia
Al Albania Eg Egypt Li Libya Si Sicily
An Asiatic Turkey Ga France LS Lebanon and Syria Sn Sinai
Bl Balearic Islands Gr Greece Lu Portugal Tn Tunisia
Bu Bulgaria Hs Spain Ma Morocco Tu Turkey in Europe
Range distribution maps
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Range maps of the Mediterranean basin for Range maps of Cyprus for all endemic species.
all species. Topography is shown with con- Topography is shown with contours at 100 m
tours at 1000 and 2000 m above sea level. above sea level. These maps also show the
Black dots indicate the country in which the eight botanical divisions (sensu Meikle 1979).
species is found. Red square indications Troodos Peak (eleva-
tion 1952 m).
XII