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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: alancrivellaro@gmail.com

Prof. Dr. Fritz Hans Schweingruber


Institute for Forest, Snow and Landscape Research – WSL
Zürcherstrasse 111
8903 Birmensdorf, Switzerland
email: fritz.schweingruber@wsl.ch

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

© Springer-Verlag Berlin Heidelberg 2013


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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

Modified from: ww2.bgbm.org/mcl/map.asp


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

RT RT

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
1. Introduction

Every anatomical atlas covers a defined group Wooden specimens are often associated with
of plants or a specific region. The idea of bark, mainly in archeological context. Bark
producing this reference book arose from the has been used in many cultures for food, as
lack of an extensive descriptions of the wood a source of fibers for textiles and cordage,
structure of trees and shrubs indigenous to in medicine, as writing material, die, etc.
the Eastern Mediterranean, and the continu- (Sandved et al. 1993). Bark anatomy is still
ing demand for wood identification in that rarely presented systematically for a great
region (Lev-Yadun 2008). The primary aim of number of species (Esau 1969, Roth 1981,
this atlas is to present the range of anatomi- Schweingruber et al. 2011, Zahur 1959).
cal features occurring in lignified plants from The anatomy of pith has been neglected. In
the Eastern Mediterranean, in order to facili- fact, there is no systematic description of
tate plant material identification. Secondly, the anatomy of the pith region for trees and
it is intended to create a basis for ecological shrubs from any woody floras of the world.
anatomy, comparative ecophyletic and ar- Here we describe in detail the twig’s pith and
chaeological studies. Here we present a wide bark anatomies by following a classification
range of taxonomically important anatomi- for anatomical features of those structures.
cal features occurring in plants from Cyprus, Having bark and pith anatomical descriptions
including various habits from trees, shrubs, addresses new possibilities in plant identi-
dwarf shrubs to some perennial herbs and fication. Moreover, the association of bark
woody lianas. Anatomical features of stem xy- and pith anatomical features to ecological
lem, twig bark and pith have been described and biological factors, will possibly open new
for 264 species (12 Gymnosperms, 2 Gneta- fields of ecophyletic and ecophysiological
les, 6 Angiosperms Monocotyledons and 244 investigation.
Angiosperms Dicotyledons) belonging to 71 Cyprus, which is the third largest island of
families. the Mediterranean Sea, offers a great range
Since former times, wood and timbers played of habitat types and a large number of typi-
an important role in the Mediterranean re- cally Eastern Mediterranean species, includ-
gion (Meiggs 1982). Therefore, identifying ing many endemics. On Cyprus grows about
plant species from archeobotanical material, 50 endemic woody species: for almost all of
historical objects, and remains belonging them no anatomical descriptions are avail-
to the human cultural heritage is important. able. Additionally, the indigenous flora of
Identifying such material is often difficult Cyprus is taxonomically diverse due to the
because wood anatomical features are only variety of ecological conditions found on the
partially present in plant remains. Moreover, island and because of its close geographical
wood of a few shrubs and dwarf shrubs, as position to Europe, the Middle East and north-
well as wood from not commonly used trees, ern Africa. We chose to extensively sample on
has been investigated in detail before. While Cyprus, considering that the trees and shrubs
some wood atlases for the east Mediterranean of the island are representative of the whole
region, and for some nearby areas, have been vegetation throughout the Eastern Mediterra-
published (Chudnoff 1956, Edlmann et al. nean region. Almost all woody species native
1994, Fahn et al. 1986, Grosser 1977, Huber to the island were included, some introduced
and Rouschal 1954, Niloufari 1961, Parsa (e.g. Eucalyptus) or cultivated (Citrus spp.
Pajouh and Schweingruber 1988, Saya 1957, and Prunus spp.) or ornamental species (e.g.
1959, Schmidt 1941, Schweingruber 1978, Lantana camara) as well as some species
1990) these descriptions are of limited value which have their distribution limits close to
for identification, since they are dispersed in Cyprus. For genera with a distribution range
a number of publications, and most of them that extends over the Eastern Mediterranean
were published prior to the introduction of region (e.g. Acer spp., Populus spp.), we
the IAWA codified anatomical feature lists. described only the species which grows on
Recently Akkemik and Yaman (2012) pub- Cyprus.
lished a valuable wood anatomy atlas of This book differs from traditional wood atlas-
Eastern Mediterranean species, mainly from es in some aspects which can be summarized
Turkey. as the follow:
A. Crivellaro, F. H. Schweingruber, Atlas of Wood, Bark and Pith Anatomy of Eastern Mediterranean Trees and Shrubs,
DOI 10.1007/978-3-642-37235-3_1, © Springer-Verlag Berlin Heidelberg 2013
1
- all anatomical features are presented in col- region as ornamental plants. The key objec-
our photomicrographs from double-stained tive of identification of groups of species by
sections. Sometimes microphotographs prove using a few selected unambiguous and clearly
to be more informative than the brief descrip- visible features. The use of the key in con-
tions removing most of the ambiguity that junction with the photomicrographs should
feature definitions alone would provide; make it possible to identify almost any plant
- the anatomy of twigs is presented systemati- occurring in the eastern Mediterranean on the
cally for each species describing details for base of its stem anatomy.
transverse sections of bark and pith; It should be emphasized that the number
- the anatomy of twig wood is described in of samples per species studied was usually
relation to its anatomical differences to stem restricted to one for both the stem and twig;
wood. only in few cases were two or three samples
The species are arranged in alphabetical or- cut from different plants. We are aware of
der of genus and species, within each of the the risks for the identification that may result
Gymnosperms, Gnetales, Angiosperm Mono- from our sampling strategy. With the present
cotyledons, and Angiosperm Dicotyledons. material it is not possible to demonstrate the
Each species has a coded description with species-specific anatomical variability.
IAWA softwood or hardwood lists of micro- For references on specific families or species
scopic feature numbers (Wheeler et al. 1989, refer to Gregory (1994). It is also not intend-
Richter et al. 2004). For Dicotyledonous An- ed to give a reference for the all work done
giosperm species, some new codified features on dendroecology in the Mediterranean re-
proposed by Schweingruber et al. (2011) were gion, for that refer to Cherubini et al. (2003),
also applied. Anatomical descriptions of stem and Grissino-Mayer website (Grissino-Mayer
wood are provided for transverse, radial and 2013).
tangential sections to facilitate the identifi-
cation process. For bark and pith codified
descriptions, new anatomical features lists
were developed, which are explained in the
chapter Definition of Anatomical Features.
Bark descriptions were difficult in many cases
because of a lack of a consistent internation-
ally accepted list of bark anatomical features.
Problems occurred on the classification of the
bark of Chenopodiaceae because the bark for-
mation mode of plants with successive cam-
bia seems to be different from those without
included phloem. Sieve tubes, parenchyma
cells, phelloderm and phellogen are difficult
to distinguish.
In general, anatomical descriptions are brief,
and details of little importance for identifica-
tion, or barely visible under an optical micro-
scope, have been omitted.
Every species included in this atlas is accom-
panied by a short plant description, providing
information on growth form, plant height and
habit, flower, leaf or fruit characters, general
distribution and habitat. Accompanying each
plant description is a Mediterranean basin
map or a Cyprus map for endemic species.
Species distribution maps are not provided
for species that belong to exotic floras and
that were introduced to the Mediterranean

2
2. Materials and Methods

Origin of the material studied height, distance from the soil level and stem
sampling point, and distance from the apex
The extensive sampling of plant material was and twig sampling point.
carried out on Cyprus in 2009 and 2010. We
used the range descriptions from the stand- Plant material preparation
ard references of Cyprus’ flora (Meikle 1979,
1985; Tsintides et al. 2002) to choose a From each 5-8 cm long sample a 1 cm long
sampling areas that was representative of the disk was cut from the central part, and split
geographic distribution and habitat range for longitudinally. Stem disks more than 1,5
each species. At each selected site we sam- cm in diameter were split to obtain a 1 cm³
pled one or two undisturbed individuals. sample, and when possible we kept the rays
Sampling sites on Cyprus are indicated on the running parallel to one side of the transverse
following map. section. In very small samples, the transverse
section was cut first. For twig samples special
care was taken to preserve the bark and the
pith on the same section. Tangential sections
were cut from the outermost growth ring,
from intermediate and latewood.
The sections were stained with astra blue and
safranin, dehydrated with alcohol and xylene,
and mounted in Canada balsam (Chaffey
2002; Schweingruber 2007). The cell walls
richer in cellulose stained blue and those
richer in lignin stained red. Parenchyma cells
RT were usually stained blue and the remaining
cells were generally stained red, making it
Sampling sites on Cyprus in black dots.
easier to interpret wood patterns in cross-
The red square indicates the city of Nicosia. sections.
A transmission-light microscope was used to
observe slides. Polarized filters were applied
for observing crystal presence, and for exam-
Rare or protected species were collected in ining specific cell wall features. Sections were
botanical gardens, especially from the arbore- imaged using a digital camera mounted on
tum in Athalassa National Forest Park south- the microscope. Magnifications are indicated
east from Nicosia, the capital city of Cyprus. in μm (0,001 mm) above a black scale bar in
Identification of floral or fruiting material was each picture.
carried out by the taxonomists of the Depart-
ment of Forest of the Ministry of Agriculture, Wood density
Natural Resources and Environment of Cy-
prus, coauthors of this book. For each plant Wood density values were calculated for each
two samples were taken: one from the main of the species studied. Wood density was
stem and another one from a twig. The stem measured on stem sample portions after
portion was near the plant’s base, and the the bark and the pith were removed. Fresh
twig portion was from the top of the stem, or wood volume was measured using Archime-
from a branch, selecting 2-4 year old shoot des’ principal by immersing the segment
portions and avoiding short shoots. The stem in a beaker containing distilled water on a
and twig samples were stored in a sealed balance. Once fresh wood volume was meas-
plastic bag to which several drops of 40% eth- ured, the sample was dried at 103±2°C for
anol were added and kept at 3-4°C until they 48 hours, allowing us to determine the wood
were sectioned. Each plastic bag was identi- mass. Wood density was calculated on the
fied with a preprinted label that included the mass/fresh volume ratio basis (g/cm³).
plant name, site location and elevation, plant

A. Crivellaro, F. H. Schweingruber, Atlas of Wood, Bark and Pith Anatomy of Eastern Mediterranean Trees and Shrubs,
DOI 10.1007/978-3-642-37235-3_2, © Springer-Verlag Berlin Heidelberg 2013
3
3. Definition of Anatomical Features

Xylem cal features” proposed by Schweingruber et al.


(2011). At the same time some IAWA features
The anatomy of gymnosperms’ xylem was de- were not considered because of their uncer-
scribed following the IAWA list of microscopic tain determination in the observed samples.
features for softwood identification (Richter et Features not applied for coding described spe-
al. 2004). cies have been omitted.
Xylem anatomy of angiosperm dicotyledon- The next pages give a short description of all
ous was described on the basis of the IAWA the xylem anatomical features detected in the
list of microscopic features for hardwood material studied. Where features were already
identification (Wheeler et al. 1989). As this described in the IAWA list (Wheeler et al.
list was proposed for tree and shrub xylems, 1989), we use their descriptions below. Where
it was slightly modified to fit with the ana- we have added features that were not included
tomical features observed in the collected in the IAWA hardwood list, we used the nota-
samples. In particular a number of comple- tion in the form “X.#” where “X” is the modi-
mentary features or character states were fied relative IAWA character number and “#” is
inserted following the “Definition of anatomi- a consecutive number.

Growth rings distinctness


1 - Growth ring boundaries distinct
Growth rings with an abrupt structural
change at the boundaries between
them. Growth rings number can be eas-
ily and exactly determined.

Rosa micrantha (100x) Arbutus unedo (100x)


1.1 - Growth ring boundaries distinct
only along some radii
The structural change at the boundaries
between growth rings is visible only
along some radii or only in between
some parenchyma rays.

Astragalus echinus (200x) Rubus sanctus (200x)


2 - Growth ring boundaries indistinct
or absent
Growth ring boundaries marked by
more or less gradual structural changes
at their poorly defined boundaries, or
not visible. Growth ring number cannot
be easily determined and their number
is uncertain.

Hedysarum cyprium (100x) Suaeda aegyptica (400x)


A. Crivellaro, F. H. Schweingruber, Atlas of Wood, Bark and Pith Anatomy of Eastern Mediterranean Trees and Shrubs,
DOI 10.1007/978-3-642-37235-3_3, © Springer-Verlag Berlin Heidelberg 2013
5
2.1 - Only one ring in the observed
sample

This feature was applied for annual


plants, for perennial specimens that
did not show any annual growth ring
boundary, and for 1-year-old twig sam-
ples.

Origanum majorana (100x) Phytolacca pruinosa (100x)


Wood porosity
3 - Wood ring-porous
Wood in which the earlywood vessels
are distinctly larger than those in the
latewood of the previous and of the
same growth ring, and form a well-de-
fined zone or ring, and in which there
is an abrupt transition from earlywood
to latewood of the same growth ring.

Rosa canina (100x) Berberis cretica (100x)


4 - Wood semi-ring-porous
Wood in which the vessels in the early-
wood are distinctly larger than those
in the latewood of the previous growth
ring, but in which there is a gradual di-
ameter decrease from large earlywood
vessels to narrower latewood vessels
of the same growth ring.

Tamarix tetrandra (100x) Phlomis cypria (200x)

5 - Wood diffuse-porous
Wood in which the vessels have more
or less the same diameter throughout
the entire growth ring. This category
also includes woods in which the lat-
est formed vessels in the latewood
are considerably smaller than those of
the earlywood of the next ring, but in
which the vessel diameter is uniform
throughout most of the growth ring.
Pyrus communis (100x) Quercus coccifera (40x)

Vessel arrangement
6 - Vessels in tangential bands
Vessels arranged perpendicularly to
rays and forming discontinuous short
or continuous tangential bands. These
bands can be straight or wavy.

6 Sambucus nigra (40x) Ulmus canescens (40x)


7 - Vessels in diagonal and/or radial
pattern

Vessels arrangement intermediates be-


tween tangential and radial (dendritic).
Transition between diagonal and den-
dritic distribution may occur within an
individual. In ring-porous woods, only
the intermediate wood and latewood
was examined.

Cercis siliquastrum (40x) Anagyris foetida (40x)


8 - Vessels in dendritic pattern

Vessels arranged in a branching radial


pattern, forming distinct zones, sepa-
rate by areas without vessels.

Rosmarinus officinalis (40x) Rhamnus alaternus (100x)

Solitary vessels
9 - Vessels predominantly solitary
Most of the vessels appear not to con-
tact another vessel. Vessels are in pairs
only by chance, most apparent pairs
actually are overlapping ends of vessel
elements as seen in transverse sections.

Vitex agnus-castus (40x) Pistacia atlantica (40x)


Vessel groupings
9.1 - Vessels in short radial multiples

Commonly radial files of 2 to 4 adjacent


vessels.

Acacia saligna (40x) Dodonaea viscosa (40x)


10 - Vessels in radial multiples of 4 or
more common

Commonly radial rows of 4 or more


adjacent vessels.

Nicotiana glauca (40x) Thevetia neriifolia (200x) 7


11 - Vessel clusters common
Groups of 3 or more vessels having
both radial and tangential contacts.
Groups about as wide radially as tan-
gentially.

Ruta chalepensis (100x) Schinus molle (100x)


Perforation plates
13 - Simple perforation plates
Perforation plates with a single circular
or elliptical opening.

Salvia fruticosa (200x) Plocama calabrica (400x)

14 - Scalariform perforation plates


Perforation plates with elongated and
parallel openings separate by one to
many mainly unbranched bars.

Fagus orientalis (400x) Platanus orientalis (400x)

15 - Scalariform perforation plates


with less than 10 bars

16 - Scalariform perforation plates


with 10-20 bars

17 - Scalariform perforation plates


with 20-40 bars

18 - Scalariform perforation plates


with more than 40 bars
Styrax officinalis (400x) Viburnum tinus (400x)

19 - Foraminate perforation plates


Perforation plates with circular or el-
liptical openings like a sieve.

8 Ephedra foeminea (400x) Ephedra nebrodensis (400x)


Intervessel pit arrangement
20 - Intervessel pits scalariform
Horizontally elongated or linear in-
tervessel pits arranged in a ladder-like
series.

Vitis vinifera (100x) Smilax aspera (400x)


21 - Intervessel pits opposite
Intervessel pits arranged in short to
long horizontal rows across the length
of the vessel.

Platanus orientalis (400x) Alnus orientalis (400x)

22 - Intervessel pits alternate


Intervessel pits arranged in diagonal
rows.

Punica granatum (400x) Capparis spinosa (400x)

Intervessel pit diameter


24 - Intervessel pits minute (< 4 μm)

25 - Intervessel pits small (4-7 μm)

26 - Intervessel pits medium (7-10 μm)

27 - Intervessel pits large (> 10 μm)

Lantana camara (400x) Juglans regia (400x)


Vessel-ray pitting
30 - Vessel-ray pits with distinct bor-
ders, similar to intervessel pits in
size and shape throughout the ray
cell
31 - Vessel-ray pits rounded or angular
with large apertures
32 - Vessel-ray pits with large
horizontal/vertical apertures
Pistacia terebinthus (400x) Quercus coccifera (400x) 9
Vessel helical thickenings
36 - Helical thickenings in vessel ele-
ments present

Ridges on the inner face of the vessel


element in a roughly helical pattern.
Very thin and thick helical thickenings
were marked and described.

Arbutus unedo (200x) Hippocrepis emerus (400x)


Vessels cell wall thick
39.1 - Vessels cell wall thickness more
than 2 μm
This feature is well seen if the vessels
cell wall is thicker than surrounding
cell tissues.

Astragalus echinus (400x) Dodonaea viscosa (400x)

Tangential diameter of vessel lumen

40.1 - Mean tangential diameter of


earlywood vessel lumina less
than 20 μm

Scabiosa cyprica (100x) Silene fruticosa (100x)

40.2 - Mean tangential diameter of


earlywood vessel lumina
20-50 μm

Artemisia arborescens (100x) Syringa vulgaris (100x)

41 - Mean tangential diameter of


earlywood vessel lumina
50-100 μm

10 Alhagi maurorum (100x) Alhagi graegorum (100x)


42 - Mean tangential diameter of early-
wood vessel lumina 100-200 μm

Cyprinia gracilis (100x) Robinia pseudoacacia (100x)


43 - Mean tangential diameter of early-
wood vessel lumina more than 200
μm

Vitis vinifera (100x) Ailanthus altissima (100x)

Vessels dimorphism
45 - Vessels of two distinct diameter
classes, wood not ring-porous
Wood with a distinct bimodal distribu-
tion of tangential diameters of vessel
lumina. Feature not applied in ring-
porous wood.

Capparis spinosa (100x) Genista sphacelata (100x)

Vessels per mm2


46 - Less than 5 earlywood vessels per
mm2

47 - 5-20 vessels per mm2

Ficus sycomorus (100x) Quercus ilex (100x)

48 - 20-40 vessels per mm2

49 - 40-100 vessels per mm2

Calicotome villosa (100x) Cistus creticus (100x) 11


50.1 - 100-200 earlywood vessels per
mm2

50.2 - More than 200 earlywood


vessels per mm2

Myrtus communis (100x) Cydonia oblonga (100x)


Vessel elements length
52.1 - Earlywood vessel element length
less than 50 μm

52.2 - Earlywood vessel element length


50-100 μm

52.3 - Earlywood vessel element length


100-200 μm

Morus alba (200x) Origanum majorana (400x)

53.1 - Earlywood vessel element length


200-500 μm

53.2 - Earlywood vessel element length


more than 500 μm

Syringa vulgaris (200x) Prunus avium (100x)


Tyloses and deposits
56 - Tyloses common
Outgrowths from an adjacent ray or
axial parenchyma cell through a pit in
a vessel wall, partially or completely
blocking the vessel lumen, and of com-
mon occurrence.

Morus alba (100x) Pistacia terebinthus (100x)


58 - Gums and other deposits in heart
wood vessels

A wide range of substances in vessels


lumen was noted.

12 Tamarix tetragyna (100x) Onosis spinosa (200x)


Imperforate tracheary elements
60 - Vascular/vasicentric tracheids present
Imperforate cells resembling in size,
shape, pitting, and wall ornamentation
narrow vessel elements and intergrad-
ing with the latter. Also including im-
perforate cells with numerous distinctly
bordered pits in their radial and tangen-
tial walls, present around the vessels,
and different from ground tissue fibers.
Pistacia terebinthus (400x) Glycyrrhiza glabra (400x)

Ground tissue fiber pits


61 - Fibers with simple to minutely
bordered pits
Fibers (libriform fibers) with simple pits
or bordered pits with the chambers less
than 3 μm in diameter.

Juglans regia (400x) Citrus sinensis (400x)

62 - Fibers with distinctly bordered pits


Fibers (or fiber-tracheids or ground tis-
sue tracheids) with bordered pits with
chambers over 3 μm in diameter.

Rubus sanctus (400x) Lomelosia cyprica (400x)

Fiber helical thickenings


64 - Helical thickenings in ground tissue
fibers
Helical thickenings in ground tissue
fibers, or ridges on the inner face of the
fiber wall in a roughly helical pattern.

Brassica hilarionis (400x)


Septate fibers
65 - Septate fibers present
Fibers with thin, unpitted, transverse
wall(s).

Lithodora hispidula (200x) Vitis vinifera (200x) 13


Fiber bands
67 - Parenchyma-like fiber bands
alternating with ordinary fibers
Tangential bands of relatively thin-
walled fibers alternating with bands of
thicker-walled fibers.

Acer pseudoplatanus (100x) Alhagi maurorum (200x)

Fiber wall thickness


68 - Fibers very thin-walled
Fiber lumina 3 or more times wider
than the double wall thickness.

Alnus orientalis (100x) Nicotiana glauca (400x)

69 - Fibers thin- to thick-walled


Fiber lumina less than 3 times the dou-
ble wall thickness, and distinctly open.

Arbutus unedo (400x) Acer obtusifolium (400x)

70 - Fibers very thick-walled


Fiber lumina almost completely closed.

Prunus dulcis (400x) Syringa vulgaris (400x)


Tension wood
70.2 - Tension wood present
The gelatinous layer, blue-stained, is
visible in fibers. Gelatinous layers usu-
ally separate from the other cell wall
layers during sectioning.

14 Acer pseudoplatanus (200x) Salix alba (100x)


Flat marginal fibers
70.3 - Radial flat marginal fibers
present
Thick-walled fibers that form a more or
less continuous layer of variable width
at the margins of a growth ring.

Scabiosa cyprica (400x) Lonicera etrusca (400x)

Axial parenchyma
75 - Axial parenchyma absent,
extremely rare, or not recognizable

Teucrium cypricum (200x) Micromeria myrtifolia (200x)

Apotracheal axial parenchyma


76 - Axial parenchyma diffuse
Single parenchyma strands or pairs of
strands distributed irregularly among
the fibrous elements of the wood.

Corylus avellana (200x) Elaeagnus angustifolia (200x)

77 - Axial parenchyma diffuse in


aggregates
Parenchyma strands grouped into short
discontinuous tangential or oblique
lines.

Rhus coriaria (100x) Ceratonia siliqua (100x)


Paratracheal axial parenchyma
78 - Axial parenchyma scanty parat-
racheal
Occasional parenchyma cells associ-
ated with the vessels, usually related
to paratracheal diffuse parenchyma,
which are immediately adjacent to ves-
sels.

Myrtus communis (100x) Prunus avium (200x) 15


79 - Axial parenchyma vasicentric
Parenchyma cells forming a complete
sheath of parenchyma around a soli-
tary vessel or vessel multiple.

Zyzyphus lotus (100x) Tamarix smyrnensis (200x)


79.1 - Axial parenchyma pervasive
The ground tissue consist exclusively
of thin-walled, unlignified parenchyma.

Frankenia hirsuta (200x) Silene galataea (200x)

83 - Axial parenchyma confluent


Coalescing vasicentric or aliform pa-
renchyma surrounding or to one side
of two or more vessels, and often
forming irregular bands.

Ceratonia siliqua (40x) Prosopis farcta (100x)

84 - Axial parenchyma unilateral


paratracheal

Paratracheal parenchyma forming


semi-circular hoods or caps only on
one side of the vessels and which can
extend tangentially or obliquely in an
aliform or confluent or banded pattern.

Cyprinia gracilis (100x) Melaleuca armillaris (100x)

Banded parenchyma
85 - Axial parenchyma bands more
than three cells wide

16 Ficus carica (100x) Acacia saligna (100x)


86 - Axial parenchyma in narrow
bands or lines up to three cells
wide

Juglans regia (100x) Diospiros kaki (100x)


89 - Axial parenchyma in marginal or in
seemingly marginal bands

Parenchymatous bands that form a


more or less continuous layer of vari-
able width at the margins of a growth
ring.

Convolvulus oleifolius (200x) Achillea cretica (100x)

89.1 - Parenchyma marginal thin-walled,


dark in polarized light

Convolvulus oleifolius (200x) Cichorium spinosum (200x)

89.2 - Ring shake


Thin-walled parenchyma cells rings that
typically break along the ring bounda-
ries during sample preparation proce-
dures.

Arabis purpurea (100x) Alyssum cypricum (400x)

Ray width
96 - Rays exclusively uniseriate

Populus nigra (200x) Calluna vulgaris (400x) 17


96.1 - Rays predominantly uniseriate
90% or more rays are uniseriate, the re-
maining rays are usually 1 to 3 seriate.

Convolvulus oleifolius (100x) Thymelaea tartonraira (100x)


97 - Ray width mostly 1-3 cells

Styrax officinalis (100x) Citrus limon (100x)

98 - Larger rays commonly 4-10


seriate

Morus nigra (100x) Brassica hilarionis (100x)

99 - Larger rays commonly more than


10 seriate

Berberis cretica (40x) Rosa micrantha (40x)

Stem lobed
99.2 - Stem lobed
The external shape of the stem in
transverse section is not roundish, but
is divided in multiple parts by radial
fissures of various depth.

18 Fumana thymifolia (4x) Teucrium micropodioides (40x)


Rays height
100 - Rays with multiseriate portion(s)
as wide as uniseriate portions

Ficus carica (100x) Diospiros kaki (400x)


100.1 - Rays confluent with ground tissue
Lateral border of ray (tangential section)
merges with axial tissue.

Salvia lanigera (200x) Lycium schweinfurthii (200x)

Rays not visible in polarized light


100.2 - Rays not visible in polarized
light

Astragalus echinus (100x) Alyssum chondrogynum (100x)

Aggregate rays
101 - Aggregate rays
Lateral border of ray (tangential section)
merges with axial tissue.

Quercus alnifolia (40x) Alnus orientalis (200x)

Rays height
102 - Ray height > 1 mm
The large rays commonly exceeding
1 mm high.

Rosa canina (40x) Hedela helix (40x) 19


Rays of two distinct sizes
103 - Rays of two distinct sizes
When viewed in tangential section, rays
form two distinct groups by their width
and usually also by their height.

Ficus sycomorus (40x) Erica sicula (100x)


Rays cellular composition
104 - All ray cells procumbent

Robinia pseudoacacia (100x) Quercus ilex (100x)

105 - All ray cells upright and/or square

Plocama calabrica (200x) Lavatera bryoniifolia (100x)

106 - Body ray cells procumbent with


one row of upright and/or
square marginal cells

Pistacia lentiscus (200x) Anagyris foetida (100x)

107 - Body ray cells procumbent with


mostly 2-4 rows of upright and/or
square marginal cells

20 Sambucus nigra (100x) Prunus domestica (100x)


108 - Body ray cells procumbent with
over 4 rows of upright and/or
square marginal cells

Diospiros kaki (100x) Morus alba (100x)


109 - Rays with procumbent, square
and upright cells mixed through-
out the ray

Laurus nobilis (100x) Prunus armeniaca (200x)

Rays per mm
1145D\VSHUPP”PP

115 - Rays per mm 4-12/mm

Berberis cretica (100x) Ficus carica (100x)

116.1 - Rays per mm 12-20/mm

116.2 5D\VSHUPP•PP

Phlomis cypria (100x) Pterocephalus multiflorus (100x)

Wood raylessness
117 - Wood rayless
Wood with only axial elements.

Salicornia fruticosa (400x) Onosma mitis (100x) 21


Storied structure
119 - Low rays storied, high rays not
storied and axial parenchyma
and/or vessel elements storied

Genista fasselata (100x) Diospiros kaki (100x)


120 - Axial parenchyma and/or vessel
elements storied

Tamarix smyrnensis (200x) Robinia pseudoacacia (200x)

121 - Fibers storied

Tamarix smyrnensis (40x) Onosis spinosa (100x)

Oil cells
124 - Oil and/or mucilage cells
associated with ray parenchyma

Laurus nobilis (200x) Vitis vinifera (200x)

Radial canals
130 - Radial canals
Tubular intercellular duct surrounded
by an epithelium present in rays.

22 Pistacia atlantica (400x) Schinus terebinthifolius (400x)


Included phloem
133.1 - Concentrically arranged single
vascular bundles

133.2 - Concentric continuous

Noaea mucronata (200x) Arabis purpurea (200x)


Prismatic crystals
136 - Prismatic crystals present
Solitary rhombohedral or octahedral
crystals, which are birefringent under
polarized light.

Punica granatum (200x) Acacia saligna (400x)

136.1 - Prismatic crystals in ray


parenchyma cells

141.1 - Prismatic crystals in axial


parenchyma cells

Ziziphus lotus (200x) Punica granatum (200x)

Crystal druses
144 - Druses present

145 - Druses in ray parenchyma cells

146 - Druses in axial parenchyma cells

Prunus domestica (400x) Prunus dulcis (200x)

Other crystals
149 - Raphides

151 - Styloids and/or elongate crystals

Laurus nobilis (400x) 23


Bark
The term ‘bark’ includes all the tissues outside
the vascular cambium (Trockenbrodt 1990).
In a plant also having secondary tissues, bark
includes the secondary phloem, the primary tis-
sues (cortex) that may be still present outside
the secondary phloem, the periderm, and the
dead tissues outside the periderm (Evert 2006).
Bark anatomical features have been described
in a number of papers and plant anatomy books
(Baas 2001, Carlquist 1992, Chang 1954, Evert
2006, Howard 1977, Junikka 1994, Lev-Yadun
1991, Machado et al. 2005, Manwiller 1972, Met-
calfe & Chalk 1950, Miles & Smith 2009, Trock-
enbrodt 1990, Zahur 1959). We have followed
the available literature to determine the follow-
ing codified list of features used in bark anatomy
descriptions. We used this codified list for both
Gymnosperms and Angiosperms.

Sieve tubes and sieve cells


B1 - Groups of three or more sieve tubes
or sieve cells

Groups of small sieve tubes or sieve


cells can be recognized among other
phloem cells of the axial system.

Bosea cypria (200x) Artemisia arborescens (400x)


B2 - Sieve tube or sieve cells arrange-
ment in tangential rows

Tangentially distributed sieve tube or


sieve cell groups can be recognized
among other phloem cells of the axial
system.

Fagonica cretica (400x) Caria pecan (100x)


B3 - Sieve tube or sieve cell arrange-
ment in radial rows
Sieve tubes or sieve cells arranged in
more or less long, continuous radial
rows.

24 Phytolacca pruinosa (200x) Nicotiana glauca (200x)


B4 - Collapsed sieve tubes or sieve cells
present
Sieve tubes or sieve cells more or less
collapsed or obliterated in older parts
of the phloem.

Ziziphus spina-christi (400x) Onosis spinosa (400x)


Ray dilatation
B5 - Distinct ray dilatation
All rays become dilated. Dilatation can
occur either by a tangential increase in
the number of ray cells, by expansion
of ray cells, or both.

Prosopis farcta (200x) Astragalus echinus (40x)

B6 - Only some rays become dilated


Dilatation is more frequent in larger
rays.

Rosa damascena (100x) Acacia saligna (200x)

Bark cellular composition


B7 - Sclerenchyma cells present in both
the phloem and cortex
Both fibers and sclereids present in the
phloem and cortex.

Robinia pseudoacacia (200x) Eucaliptus gomphocephala (100x)

B8 - Phloem uniform
Sclerenchyma cells absent. Phloem
composed only of sieve tubes and pa-
renchyma cells

Lomelosia cyprica (100x) Arbutus andrachne (100x) 25


Fibers
B9 - Fibers present
Fibers more or less thick-walled, lig-
nified and elongated. In the primary
phloem, fibers usually occur in the
outermost part of the tissue. In the
secondary phloem, fibers are distrib-
uted in various patterns among other
phloem cells of the axial system.

Erica carnea (100x) Plumbago europaea (400x)

B10 - Fibers with an unlignified inner-


most layer

The innermost fiber layer unlignified,


with a structure similar to the gelati-
nous fibers in tension wood (stained
blue with astrablue)

Nerium oleander (100x) Salix alba (400x)

B11 - Fibers in radial rows


Fibers in uniseriate or multiseriate regu-
lar or irregularly formed radial rows

Phagnalon rupestre (100x) Glycyrrhiza glabra (200x)

B12 - Fibers in tangential bands


Tangentially continuous rows of fib-
ers. They alternate with bands of sieve
tubes and parenchymatous components
of the axial system.

Syringa vulgaris (100x) Vitis vinifera (200x)

B13 - Fibers scattered or irregularly


dispersed

Single fiber cells dispersed in the phlo-


em.

26 Capparis spinosa (40x) Ruta chalepensis (100x)


B14 - Fibers grouped
Three or more fiber cells per group;
groups irregularly or tangentially dis-
tributed in the phloem.

Centaurea akamantis (100x) Lavatera bryoniifolia (100x)


Sclereids
B15 - Sclereids present
Sclereids very thick-walled, lignified
and relatively short with multilayered
secondary wall.

Tamarix tetragyna (100x) Viburnum opulus (200x)

B16 - Sclereids in radial rows

Tamarix dalmatica (100x) Fagus orientalis (100x)

B17 - Sclereids in tangential bands

Crataegus azarolus (40x) Ceratonia siliqua (100x)

B18 - Sclereids scattered or irregularly


dispersed

Olea europaea (100x) Cedrus brevifolia (40x) 27


B19 - Sclereids grouped tangentially
Sclereid groups in tangential clusters.

Brassica hilarionis (100x) Rhamnus alaternus (40x)


Crystals
B20 - Prismatic crystals present

B21 - Acicular crystals present

Celtis turnefortii (200x) Phytolacca pruinosa (400x)

B22 - Druses present

B23 - Crystal sand present

B24 - Raphides present

Silene fruticosa (400x) Taxus baccata (400x)

Secretory elements and canals


B25 - With single, irregularly dispersed
laticifers or ducts.

Euphorbia thompsonii (200x) Ficus carica (100x)

B26 - Ducts present

28 Pinus pinea (100x) Schinus terebinthifolius (400x)


B27 - Laticifers or intercellular canals

Eucalyptus camaldulensis (400x) Citrus paradisii (400x)

Cell content
B28 - Cell contents in parenchyma cells

Eucaliptus torquata (100x) Alhagi maurorum (100x)


B29 - Layered phellem
Phellem consist of alternating bands of
sclerenchyma and parenchyma tissues.

Noaea mucronata (200x) Teucrium cypricum (100x)

Phellem
B30 - Phellem not clearly distinct
The cellular composition doesn’t per-
mit clear separation between phloem
and phellem.

Arbutus unedo (40x) Cyprinia gracilis (100x)

B31 - Phellem and/or cortex homoge-


neous

Phellem distinct, consists of regularly


arranged square or rectangular cells.

Plocama calabrica (200x) Pyrus malus (400x) 29


B32 - Phellem and/or cortex heteroge-
neous

Phellem distinct, consists of irregular


shaped cells.

Berberis cretica (100x) Aristolochia sempervirens (100x)


B33 - Phellem and/or epidermis distinct
in polarized light

Hippocrepis emerus (100x) Inula critmoides (100x)

B34 - Lignified cells in phellem and/or


epidermis

Lonicera japonica (100x) Ilex aquifolium (100x)

Phelloderm
B35 - Layered phelloderm
Phelloderm consist of alternating bands
of sclerenchyma and parenchyma tis-
sues.

Melaleuca armillaris (40x) Calicotome villosa (100x)

30
Pith
Very few publications describe the pith
structure (Carlquist 1992, Metcalfe e
Chalk 1950, Piccioli 1919, Schweingru-
ber 2007).
The following list of features was de-
fined by observing the feature in the
slides, and is not based on the avail-
able literature.

Pith not visible in polarized light


P0.1 - Pith not visible in polarized light
If the sample is observed in polarized
light the pith area becomes completely
black.

Onosma caepistosa (100x) Cyprinia gracilis (100x)

Pith shape
P1 - Pith shape round
Round or roundish to oval in transverse
section.

Calycotome villosa (40x) Plocama calabrica (100x)

P1.1 - Pith shape polygonal


Pith shape with at least four straight
sides and angles: rhomboid, hexago-
nal, irregular and star shaped. Trian-
gular and pentagonal shapes are not
described in this features (see P1.2 and
P1.3).

Lantana camara (20x) Ceratonia siliqua (40x) 31


P1.2 - Pith shape triangular
Pith shape with three straight sides and
angles.

Citrus sinensis (40x) Zygophyllum album (40x)


P1.3 - Pith shape pentagonal
Pith shape with five straight sides and
angles.

Centaurea akamantis (40x) Rubus sanctus (20x)

P1.4 - Pith shape square, rectangular


Pith shape with four straight sides and
angles.

Punica granatum (40x) Origanum dubium (100x)

Medullary sheath
P2 - Medullary sheath present
A layer of thick-walled cells surrounding
the pith.

Ailanthus altissima (40x) Prunus domestica (40x)

Pith cellular composition


P3 - Heterogenous pith
Different cell types in various distribu-
tional patterns occur among parenchy-
matic cells.

32 Eucalyptus camaldulensis (40x) Populus alba (100x)


P3.1 - Sclereids present
Sclereids in the pith.

Myrtus communis (100x) Tamarix ahpylla (200x)


P3.2 - Fibers present
Fibers in the pith.

Eucalyptus gomphocephala (40x) Nicotiana glauca (200x)


P3.3 - Thick-walled parenchyma cells
present

Parenchyma cells with thick cell walls


present.

Brassica hilarionis (100x) Quercus ilex (100x)

P3.4 - Unlignified cells present


Pith completely or in part composed of
unlignified cell walls. In safranin-astra-
blue stained sections this cells appear
blue.

Convolvulus oleifolius (100x) Rhus coriaria (40x)


P4 - Cells dimorphic
Cells of different shape and size occur
throughout the pith.

Rosa canina (40x) Prasium majus (40x) 33


Pith cell contents
P5 - Cell contents present
Numerous kinds of substances visible
in cell lumen (gum, tannins, dark stain-
ing substances).

Pyrus syriaca (100x) Helianthemum stipulatum (200x)

Pith crystals
P6 - Prismatic crystals present

Diospiros kaki (100x) Bosea cypria (400x)

P6.1 - Acicular crystals present

P6.2 - Crystal druses present

Vitis vinifera (200x) Prunus dulcis (400x)

P6.3 - Raphides present

P6.4 - Crystal sand present

Nicotiana glauca (200x)


Pith canals
P7 - Laticifers or intercellular canals
Tube-like canals without border cells.

34 Prunus dulcis (40x) Ziziphus ziziphus (200x)


Pith cells pits
P9 - Pits in longitudinal cell walls
(radial section)
Simple pits present in longitudinal pith
cell walls.

P9.1 - Pits in transverse cell walls


Simple pits present in transverse pith
cell walls.
Prasium majus (400x) Alnus orientalis (400x)
P9.2 - Pits grouped
Simple pits aggregated into clusters of
two or more in longitudinal and/or in
transverse cell walls.

Atriplex halimus (400x) Viburnum tinus (400x)

P9.3 - Pits of two distinct size and/or


shape

Pits of two distinct diameters or shapes


in longitudinal and/or in transverse
cell walls.

Platanus orientalis (400x) Fagonia cretica (200x)

Vascular bundles
P10 - Vascular bundles in the pith
Vascular bundles completely embed-
ded in the pith.

Cyprinia gracilis (100x) Nerium oleander (200x)


P10.1 - Vascular bundles separate

Single Vascular bundles clearly separate


from one another by rays or fibers.

Rubus discolor (40x) Berberis cretica (100x) 35


P10.2 - Vascular bundles not distinct

Calluna vulgaris (200x) Olea europaea (200x)

P12 - Tracheary elements of metaxylem


in distinct radial rows
Radial files of two or more tangentially
adjacent primary vessels easily recog-
nizable.

Cercis siliquastrum (100x) Ostrya carpinifolia (400x)

Pith cells axial arrangement


P13 - Axial cells in regular rows
Pith cells in regularly arranged vertical
rows, observed in radial section.

Clematis vitalba (200x) Lonicera japonica (200x)

36
4. Identification Keys

How to use the keys Keys structure


The objective of the identification key is to WOOD WITHOUT VESSELS (GYMNOSPERMS)...38
separate groups of species by using a few - Without resin ducts
selected unambiguous and clearly visible fea- - With resin ducts
tures. Due to intraspecific anatomical variabil-
ity, correct identification can only be achieved WOOD WITH VESSELS
through comparison with illustrations and - With included phloem.............................39
descriptions in the main section of the atlas. - Phloem strands in tangential bands
Some species were repeated two or more - Rays absent
times in the key because of their not clear - Phloem strands scattered, isolated
structure. The page numbers given in the key - Rays present
refers to the complete anatomical description - Rays heterogenous
of the species. - Rays homogenous
Coniferous woods are listed under the sec- - Rays absent
tions “Wood without vessels” and are sepa- - Without included phloem........................40
rated by resin ducts presence or absence. - Ring-porous........................................40
Additional features are given to facilitate - Rays exclusively uniseriate.............40
identification process: axial parenchyma, ray - Rays heterogenous
tracheids (“+”, present; “-”, absent), and cross- - Rays homogenous
field pitting (S, small; L, large). - Larger rays up to 3 seriate..............40
Dicotyledons wood and Monocotyledons plant - Rays heterogenous
body are grouped together under the section - Rays homogenous
“Wood with vessels”. They are divided first by - Larger rays more than 4 seriate......41
presence or absence of included phloem. In - Rays heterogenous
those species with successive cambia, phloem - Rays homogenous
strands can be arranged in continuous tan- - Semi-ring-porous...............................42
gential bands, or in single, scattered vascular - Rays exclusively uniseriate.............42
bundles as seen in transverse section. Then - Rays heterogenous
species are then classified by raylessness and - Rays homogenous
ray type (heterogeneous or homogeneous). - Larger rays up to 3 seriate.............42
In woods without included phloem wood po- - Rays heterogenous
rosity is the first classification factor (ring-po- - Rays homogenous
rous, semi-ring-porous, and diffuse-porous). - Larger rays more than 4 seriate.....43
Within each of the tree groups raylessness - Rays heterogenous
and ray width were then applied as grouping - Rays homogenous
factors: rays are absent, exclusively uniseriate - Diffuse-porous...................................44
(90% or more), up to 3 cells wide, and more - Wood rayless..................................44
than 4 cells wide. Ray width classification - Rays exclusively uniseriate.............45
in the identification key always refers to the - Rays heterogenous
larger rays. Ray type is also specified (heter- - Rays homogenous
ogenous and homogenous). Within each of - Larger rays up to 3 seriate..............46
the resulting groups additional features are - Rays heterogenous
given for each specie in order to help identifi- - Rays homogenous
cation. Those additional features refer to per- - Larger rays more than 4 seriate......47
foration plates (13, simple; 14, scalariform; - Rays heterogenous
19, foraminate), helical thickenings (“+”, - Rays homogenous
present; “-”, absent), and vessel groupings (9,
vessels predominantly solitary; 10, vessels in
radial multiples; 11, vessel clusters common).

A. Crivellaro, F. H. Schweingruber, Atlas of Wood, Bark and Pith Anatomy of Eastern Mediterranean Trees and Shrubs,
DOI 10.1007/978-3-642-37235-3_4, © Springer-Verlag Berlin Heidelberg 2013
37
Wood without vessels (Gymnosperms)
The major mass of conifers wood consists of tracheids. Growth ring boundaries are usually
distinct. Axial parenchyma is present in some species. Axial and radial resin canals are pre-
sent or absent. Rays are mostly uniseriate.

Wood of a conifer without resin ducts. Wood of a conifer with resin ducts.

Cross-field pitting
Axial parenchyma
Ray tracheids

Species pg.

Without resin ducts + - S Cupressus sempervirens..............................54


+ - S Juniperus excelsa........................................56
+ - S Juniperus foetidissima.................................58
+ - S Juniperus oxycedrus....................................60
+ - S Juniperus phoenicea....................................62
- - S Taxus baccata.............................................76
- + S Cedrus brevifolia.........................................64
- + S Cedrus libani...............................................66

With resin ducts - + S Pinus brutia.................................................68


- + L Pinus nigra..................................................72
- + S Pinus halepensis..........................................70
- + S Pinus pinea..................................................74

38
Wood with vessels and with included phloem
Species with successive cambia are separated by phloem strands arranged in continuous tan-
gential bands, or in single, scattered vascular bundles as seen in transverse section.
We are aware about the great anatomical variability within plants with successive cambia.

Successive cambia with phloem strands in tangential bands. Single vascular bunbles surrounded by parenchyma cells.

Helical thickenings
Perforation plates

Vessels grouping

Species pg.

Phloem strands in tangential bands


Rays absent 13 - 11 Atriplex semibaccata.................................214
13 - 11 Salicornia fruticosa...................................220
13 - 9 Sarcocornia perennis................................222

Phloem strands scattered and/or isolated


Rays present
Rays heterogeneous 13 - 9 Convolvulus dorycnium.............................248
13 - 9 Bosea cypria.............................................100

Rays homogeneous 13 - 10 Convolvulus oleifolius ...............250, 252, 254

Rays absent 13 - 11 Arthrocnemum macrostachyum................208


13 - 11 Atriplex halimus........................................210
13 - 11 Atriplex portulacoides...............................212
13 - 11 Noaea mucronata......................................218
13 - 10 Suaeda aegyptica......................................224
13 + 11 Halocnemum strobilaceum........................216
13 + 11 Suaeda vera..............................................226
13 + 11 Ruscus aculeatus.........................................90
13 + 9 Asparagus acutifolius.................................86
13 + 9 Asparagus stipularis...................................87
13 + 9 Arundo donax.............................................88
13 + 9 Phragmites australis...................................89
13 + 9 Smilax aspera............................................91

39
Wood with vessels and without included phloem
Wood with a great variation in type, size, form and arrangement of its cell types. Earlywood ves-
sels can be much larger than those of the latewood (ring-porous woods), or vessels diameter can
be uniform throughout the growth ring (diffuse-porous woods). Intermediate pattern is frequent
(semi-ring-porous woods). Axial parenchyma and rays may or may not be present.
Ring-porous wood. Diffure-porous wood.

Helical thickenings
Perforation plates

Vessels grouping

Species pg.

Ring-porous
Rays exclusively uniseriate
Rays heterogeneous 13 - 9 Cyprinia gracilis........................................126
13 - 9 Hypericum hircinum..................................330
13 + 9; 11 Pterocephalus multiflorus..........................262

Rays homogeneous 13 - 9 Castanea sativa.........................................312


13 - 9; 10 Satureja thymbra......................................378
13 + 9 Rubia tenuifolia.........................................512

Larger rays up to 3 seriate


Rays heterogeneous 13 - 9 Convolvulus oleifolius................................254
13 - 9 Salvia willeana..........................................376
13 - 9 Fraxinus angustifolia.................................430
13 - 9; 11 Fraxinus ornus..........................................432
13 - 11 Capparis spinosa.......................................196
13 - 11 Teucrium divaricatum...............................388
13 + 9 Acanthoprasium integrifolium...................334
13 + 9; 11 Cotinus coggygria.....................................102
13 + 9; 11 Hypericum confertum................................328
13 + 11 Pistacia atlantica.......................................104
13 + 11 Pistacia lentiscus.......................................106
13 + 11 Pistacia terebinthus...................................108
13 + 11 Rhus coriaria.............................................110

40
Helical thickenings
Perforation plates

Vessels grouping
Species pg.

(Larger rays up to 3 seriate)


(Rays heterogeneous) 13 + 11 Celtis tournefortii......................................194
13 + 11 Genista fasselata......................................298
13 + 11 Phlomis cypria subsp. cypria.....................360
13 + 11 Phlomis cypria var. occidentalis.................362
Rays homogeneous
13 - 9 Fraxinus angustifolia.................................430
13 + 9; 11 Teucrium creticum....................................384
13 + 11 Robinia pseudoacacia................................308

Larger rays more than 4 seriate


Rays heterogeneous 19 - 9 Ephedra nebrodensis...................................82
13 - 9 Aristolochia sempervirens.........................124
13 - 9 Onosis spinosa subsp. leisosperma............306
13 - 9 Quercus infectoria subsp. veneris.............324
13 - 9 Rubus sanctus...........................................500
13 - 9; 11 Rubus discolor...........................................498
13 - 9; 11 Vitis vinifera..............................................568
13 - 11 Hedera helix..............................................122
13 - 11 Astragalus echinus subsp. echinus ...........292
13 - 11 Morus nigra...............................................416
13 - 11 Clematis vitalba........................................454
13 + 9 Rosa micrantha.........................................496
13 + 9; 1 Prunus armeniaca.....................................476
13 + 9; 11 Celtis australis..........................................192
13 + 9; 11 Elaeagnus angustifolia..............................266
13 + 9; 11 Morus alba................................................414
13 + 9; 11 Prunus dulcis.............................................482
13 + 9; 11 Rosa canina...............................................492
13 + 9; 11 Rosa damascena.......................................494
13 + 9; 11 Ailanthus altissima....................................538
13 + 11 Pistacia atlantica.......................................104
13 + 11 Berberis cretica.........................................154
13 + 11 Celtis tournefortii......................................194
13 + 11 Lonicera japonica......................................200

Rays homogeneous 13 - 9 Quercus cerris..........................................318


13 - 9 Quercus infectoria subsp. veneris.............324
13 + 11 Cercis siliquastrum...................................296
13 + 11 Robinia pseudoacacia................................308
13 + 11 Ulmus canescens.......................................562

41
Helical thickenings
Perforation plates

Vessels grouping
Species pg.

Semi-ring-porous
Rays exclusively uniseriate
Rays heterogeneous 13 - 9 Lomelosia cyprica......................................258
13 - 9 Lavandula stoaechas.................................340
13 - 9 Salix alba..................................................526
13 - 9 Lycium schweinfurtii.................................452
13 - 9; 11 Cistus salvifolius.......................................236
13 - 9; 11 Fumana thymifolia....................................240
13 - 9; 11 Micromeria chionistrae.............................342
13 - 9; 11 Micromeria myrtinifolia.............................346
13 - 9; 11 Polygonum equisetiforme..........................450
13 - 9; 11 Thymelaea hirsuta....................................558
13 - 11 Micromeria microphylla.............................344
13 - 11 Asperula cypria.........................................506
13 + 9; 11 Thymelaea tartoriaria...............................560
13 + 11 Pterocephalus multiflorus subsp. multiflorus.....260

Rays homogeneous 13 - 9 Helianthemum obtusifolium......................242


13 - 9 Origanum dubium.....................................354
13 - 9 Teucrium micropodioides..........................394
13 - 9 Thymus intiger..........................................398
13 - 9 Lycium schweinfurtii.................................542
13 - 9; 11 Satureja thymbra......................................378
13 - 11 Origanum cordifolium...............................352
13 + 9 Teucrium kyreniae....................................392
13 + 9 Rubia tenuifolia.........................................512
13 + 11 Teucrium cyprium.....................................386
13 + 11 Teucrium kotschyannum...........................390

Larger rays up to 3 seriate


Rays heterogeneous 19 - 9 Ephedra foeminea.......................................80
14 + 9; 11 Corylus avellana........................................158
14 + 9; 11 Viburnum tinus...........................................98
13 - 9 Cistus creticus...........................................228
13 - 9 Cistus ladanifer.........................................230
13 - 9 Helianthemum stipulatum.........................244
13 - 9 Prasium majus..........................................366
13 - 9 Salvia willeana..........................................376
13 - 9 Callistemon lanceolatus.............................418
13 - 9 Syringa vulgaris........................................438
13 - 9 Crataegus mongyna..................................470
13 - 9 Pyracantha coccinea.................................484
13 - 9 Pyrus syriaca.............................................490
13 - 9 Citrus aurantium.......................................514
13 - 9 Origanum majorana.................................356
13 - 9 Salvia fruticosa.........................................372
13 - 9 Pyrus communis........................................486

42
Helical thickenings
Perforation plates

Vessels grouping
Species pg.

(Larger rays up to 3 seriate)


(Rays heterogeneous) 13 - 9; 11 Pyrus malus..............................................488
13 - 10 Nicotiana glauca.......................................544
13 - 11 Ptilostemon chamapeuce...........................152
13 - 11 Capparis spinosa ......................................196
13 - 11 Convolvulus dorycnium..............................248
13 - 11 Anagyris foetida........................................288
13 - 11 Teucrium divaricatum subsp. canescens....388
13 - 11 Prosopis farcta..........................................408
13 + 9 Hippocrepis emerus...................................304
13 + 9 Ziziphus spina-christi.................................462
13 + 9 Cotoneaster nummuralis...........................466
13 + 9 Crataegus azarolus...................................468
13 + 9 Cydonia oblonga.......................................474
13 + 9 Citrus limon..............................................516
13 + 9; 11 Ostrya carpinifolia....................................160
13 + 9; 11 Prunus avium............................................478
13 + 9; 11 Artemisia arborescens...............................134
13 + 9; 11 Arbutus andrachne...................................268
13 + 9; 11 Spartium junceum.....................................310
13 + 9; 11 Hypericum confertum subsp. stenobotrys...328
13 + 9; 11 Phlomis brevibracteata.............................358
13 + 9; 11 Sorbus cretica...........................................504
13 + 11 Arbutus unedo...........................................270
13 + 11 Calycotome villosa.....................................294
13 + 11 Genista fasselata......................................298
13 + 11 Lavandula angustifolia..............................338
13 + 11 Phlomis cypria subsp. cypria.....................360
13 + 11 Phlomis cypria subsp. occidentalis............362
13 + 11 Phlomis lunariifolia...................................364
13 + 11 Rosmarinus officinalis...............................368
13 + 11 Salvia dominica.........................................270

Rays homogeneous 13 - 9 Sideritis cypria..........................................382


13 - 9; 11 Odontites linkii..........................................440
13 - 9; 11 Plocama calabrica.....................................508
13 + 9; 11 Pterocephalus multiflorus subsp. obtusifolium..262
13 + 9; 11 Spartium junceum.....................................310
13 + 9; 11 Teucrium creticum....................................384

Larger rays more than 4 seriate


Rays heterogeneous 14 - 9; 11 Styrax officinalis.......................................548
13 - 9 Aristolochia sempervirens.........................124
13 - 9 Dittrichia viscosa......................................140
13 - 9 Phagnalon rupestre..................................150
13 - 9 Erica arborea............................................274

43
Helical thickenings
Perforation plates

Vessels grouping
Species pg.

(Larger rays more than 4 seriate)


(Rays heterogeneous) 13 - 9 Rubus sanctus...........................................500
13 - 9 Tamarix tetragyna....................................554
13 - 9; 11 Rubus discolor...........................................498
13 - 9; 11 Citrus sinensis...........................................518
13 - 9; 11 Tamarix smyrnensis..................................552
13 - 9; 11 Tamarix tetrandra....................................556
13 - 11 Hedera helix..............................................122
13 - 11 Hirtellina lobelii.........................................146
13 - 11 Convolvulus dorycnium.............................248
13 - 11 Argyrolobium uniflorum............................290
13 - 11 Astragalus echinus subsp. echinus...........292
13 - 11 Sarcopoterium spinosum...........................502
13 + 9 Prunus domestica......................................480
13 + 9; 11 Helichrysum italicum................................142
13 + 9; 11 Prunus dulcis............................................482
13 + 9; 11 Rosa damascena.......................................494
13 + 11 Lithodora hispidula...................................164
13 + 11 Lonicera japonica......................................200
13 + 11 Nepeta trodii.............................................350
13 + 11 Tamarix dalmatica....................................550
Larger rays more than 4 seriate
Rays homogeneous 13 - 9 Juglans regia.............................................332

Diffuse-porous
Wood rayless 13 - 9 Alyssum troodi..........................................178
13 - 9 Arabis purpurea........................................182
13 - 9 Erysimum kykkoticum...............................186
13 - 9 Silene galatea............................................206
13 - 9 Frankenia hirsuta......................................326
13 - 9; 11 Alyssum akamasicum................................172
13 - 9; 11 Dianthus cyprius.......................................202
13 - 9; 11 Onosma mitis............................................170
13 - 9; 11 Arabis cypria............................................180
13 - 9; 11 Salicornia fruticosa...................................220
13 - 9; 11 Sarcocornia perennis................................222
13 - 11 Onosma fruticosa......................................168
13 - 11 Arthrocnemum macrostachyum................208
13 - 11 Atriplex halimus........................................210
13 - 11 Atriplex portulacoides...............................212
13 - 11 Atriplex semibaccata.................................214
13 - 11 Noaea mucronata.....................................218
13 + 9 Silene fruticosa..........................................204
13 + 9 Plumbago europaea..................................448
13 + 9; 11 Suaeda vera..............................................226
13 + 11 Onosma caespitosa...................................166
13 + 11 Halocnemum strobilaceum........................216

44
Helical thickenings
Perforation plates

Vessels grouping
Species pg.

Rays exclusively uniseriate


Rays heterogeneous 14 - 9; 11 Viburnum opulus........................................96
13 - 9 Helianthemum syriacum...........................246
13 - 9 Calluna vulgaris........................................272
13 - 9 Eucalyptus torquata..................................424
13 - 9 Melaleuca armillaris..................................426
13 - 9 Ziziphus lotus............................................460
13 - 9 Salix alba..................................................526
13 - 9 Lycium schweinfurtii.................................542
13 - 9 Fagonia cretica.........................................570
13 - 9; 11 Punica granatum......................................402
13 - 9; 11 Fumana thymifolia....................................240
13 - 9; 11 Convolvulus oleifolius var. oleifolius...........252
13 - 10 Euphorbia thompsonii................................280
13 - 10; 11 Thymelaea hirsuta....................................558
13 - 11 Calamintha incana....................................336
13 - 11 Micromeria microphylla............................344
13 - 11 Thymbra capitata.....................................396
13 - 11 Asperula cypria.........................................506
13 + 10 Dodonaea viscosa.....................................534
13 + 11 Lycium ferocissimum................................540

Rays homogeneous 14 - 10; 11 Alnus orientalis.........................................156


13 - 9 Fumana arabica........................................238
13 - 9 Helianthemum obtusifolium......................242
13 - 9 Scutellaria sibthorpii.................................380
13 - 9 Teucrium micropodioides..........................394
13 - 9 Thymus intiger..........................................398
13 - 9 Lycium schweinfurtii.................................542
13 - 9; 11 Micromeria nervosa..................................348
13 - 9; 11 Salvia lanigera..........................................374
13 - 9; 11 Aesculus hippocastanum...........................532
13 - 10 Euphorbia hierosolymitan.........................278
13 - 10 Populus alba.............................................522
13 - 10 Populus nigra............................................524
13 - 11 Origanum cordifolium...............................352
13 + 9 Teucrium kyreniae....................................392
13 + 9 Plumbago europaea..................................448
13 + 9 Cydonia oblonga.......................................474
13 + 9 Rubia laurae.............................................510
13 + 9; 11 Vinca major..............................................118
13 + 11 Teucrium cyprium.....................................386

45
Helical thickenings
Perforation plates

Vessels grouping
Species pg.

Larger rays up to 3 seriate


Rays heterogeneous 14 - 9 Buxus sempervirens..................................188
14 - 9 Cornus sanguinea.....................................256
14 - 9 Laurus nobilis...........................................398
14 + 9; 11 Corylus avellana.......................................158
14 + 9; 11 Viburnum tinus...........................................98
13 - 9 Cistus creticus...........................................228
13 - 9 Cistus parvifolius......................................234
13 - 9 Helianthemum stipulatum.........................244
13 - 9 Helianthemum syriacum...........................246
13 - 9 Convolvulus oleifolius var. desertii............250
13 - 9 Diospiros kaki...........................................264
13 - 9 Ricunus communis....................................282
13 - 9 Alhagi graegorum.....................................284
13 - 9 Alhagi maurorum.....................................286
13 - 9 Laurus nobilis...........................................398
13 - 9 Ficus carica...............................................410
13 - 9 Eucalyptus gomphocephala.......................422
13 - 9 Myrtus communis......................................428
13 - 9 Ziziphus ziziphus.......................................464
13 - 9 Crataegus mongyna..................................470
13 - 9 Crataegus x sinaica...................................472
13 - 9 Pyracantha coccinea.................................484
13 - 9 Pyrus syriaca............................................490
13 - 9 Citrus aurantium......................................514
13 - 9; 11 Nerium oleander.......................................116
13 - 9; 11 Glycirrhiza glabra.....................................300
13 - 9; 11 Origanum majorana.................................356
13 - 9; 11 Oleae europaea.........................................434
13 - 9; 11 Pyrus communis........................................486
13 - 9; 11 Pyrus malus..............................................488
13 - 9; 11 Lantana camara........................................564
13 - 10 Suaeda aegyptiaca....................................224
13 - 10 Nicotiana glauca.......................................544
13 - 10; 11 Cichorium spinosum..................................138
13 - 11 Sambucus nigra..........................................94
13 - 11 Ptilostemon chamapeuce...........................152
13 - 11 Ceratonia siliqua.......................................190
13 - 11 Phillyrea latifolia.......................................436
13 - 11 Rhamnus alaternus...................................456
13 - 11 Withania somnifera...................................546
13 + 9 Cistus monspelliensis.................................232
13 + 9 Hippocrepis emerus...................................304
13 + 9 Ziziphus spina-christi................................462
13 + 9 Cotoneaster nummuralis...........................466
13 + 9 Crataegus azarolus...................................468
13 + 9 Citrus limon..............................................516

46
Helical thickenings
Perforation plates

Vessels grouping
Species pg.

(Rays heterogeneous) 13 + 9 Vitex agnus-castus....................................566


13 + 9 Zygophyllum album...................................572
13 + 9; 11 Schinus terebinthifolia...............................114
13 + 9; 11 Acer pseudoplatanus.................................530
13 + 9; 11 Artemisia arborescens..............................134
13 + 9; 11 Lonicera etrusca.......................................198
13 + 9; 11 Arbutus andrachne...................................268
13 + 9; 11 Sorbus cretica...........................................504
13 + 10 Plumbago auriculata.................................446
13 + 10 Dodonaea viscosa......................................534
13 + 10; 11 Schinus molle............................................112
13 + 11 Arbutus unedo...........................................270
13 + 11 Lavandula angustifolia..............................338
13 + 11 Salvia dominica.........................................370
13 + 11 Rhamnus lycioides.....................................458
13 + 11 Ruta chalepensis.......................................520

Rays homogeneous 13 - 9 Convolvulus oleifolius var. desertii............250


13 - 9 Eucalyptus camaldulensis..........................420
13 - 9 Phytolacca pruinosa..................................442
13 - 9; 11 Acacia saligna...........................................406
13 - 9; 11 Odontites linkii..........................................440
13 - 11 Alyssum cypricum.....................................176
13 + 9; 11 Acer pseudoplatanus.................................530

Larger rays more than 4 seriate


Rays homogeneous 13 - 9 Quercus ilex..............................................322
13 - 11 Ambrosia maritima...................................132
13 + 9 Acer obtusifolium......................................528

47
5. Anatomical Description of Species

This chapter contains anatomical


descriptions of stem and twig xylem,
as well as the bark and pith regions
of the twigs of 264 species belonging
to 71 families. Species are listed in
alphabetical order within families.
For a complete species list refer to
page 579.

Anatomical description of species

A. Crivellaro, F. H. Schweingruber, Atlas of Wood, Bark and Pith Anatomy of Eastern Mediterranean Trees and Shrubs,
DOI 10.1007/978-3-642-37235-3_5, © Springer-Verlag Berlin Heidelberg 2013
How to read species descriptions

Whole plant or plant detail photograph

Botanical name with synonyms

English common name

Plant description of habitus,


plant height, and ecological
preferences

Distribution map over the


Mediterranean basin or on
Cyprus for endemic species

Twig bark anatomical descrip-


tion

Twig xylem description in


relation to its differences to
stem wood anatomy

Twig pith anatomical descrip-


tion

Photomicrographs with scale bars


in microns (0,001 mm)

50
Codified anatomical feature descriptions
for stem wood, twig bark and pith

Description of anatomical features visible


in stem wood transverse section

Family name

Description of anatomical
features visible in stem
wood radial section

Description of anatomical
features visible in stem
wood tangential section

Botanical name

Basic density values for stem wood

51
Gymnosperms

This section contains anatomical de-


scriptions of stem and twig xylem, as
well as the bark and pith regions of
12 coniferous species belonging to 3
families.

Gymnosperms
Cupressus sempervirens L.
Mediterranean Cypress
Cupressaceae

PLANT DESCRIPTION
Evergreen, resinous tree up to 30 m
high, with broad (var. horizontalis) or
narrowly conical crown (var. sempervi-
rens). Leaves scale-like, 2–5 mm long.
Native to eastern Mediterranean and
northern Iran. Indigenous to Cyprus,
abundantly planted (0-1400 m alt.).

TWIG ANATOMY
Bark
Groups of sieve cell in radial rows.
Sclerenchyma cells in phloem and
in cortex. Fibers in single tangential
rows in phloem, scattered and irregu-
larly dispersed in cortex. Crystal sand
present, mostly in rays. Phellem ho-
mogeneous, distinct in polarized light.
Cortex consist of thin-walled unligni-
fied parenchyma cells.

Xylem
Rays exclusively uniseriate. Average
ray height very low (no more than 4
cells).
›T

Pith
Pith shape polygonal. Thick-walled
Cupressus sempervirens L.

parenchyma cells present. Cells dimor-


phic. Pits in transverse and in longi-
tudinal cell walls. Pits of two distinct
sizes. Vascular bundles not clearly
distinct.

›T

54
Stem xylem: 40 44 50 55 56 72 73 74 76 80 85 87 92 98 104 108
Twig bark: B3 B7 B9 B12 B23 B33
Pith: P1.1 P3.3 P4 P9 P9.1 P9.3 P10.2

Cupressaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct. Late-
wood tracheids thick-walled (double
wall thickness larger than radial lumen
diameter). Axial parenchyma present,
diffuse and tangentially zonate.

›T ›T

Radial section
Pitting in radial walls of earlywood
tracheids predominantly uniseriate.
Average tracheids length short (less
than 3000 μm). Axial parenchyma
transverse end walls smooth. Ray tra-
cheids absent or very rare. End walls
of ray parenchyma cells smooth. Hori-
zontal walls of ray parenchyma cells
smooth (unpitted). Cross-field pitting
piceoid. Number of pits per earlywood
cross-field in earlywood 1-3.

›T ›T

Tangential section
Average ray height high (from 16 to 30
Cupressus sempervirens L.
cells). Some rays in part 2-3 seriate.

Wood basic density: 0,47 g/cm³

›T ›T

55
Juniperus excelsa M. Bieb. subsp. excelsa
Grecian Juniper
Cupressaceae

PLANT DESCRIPTION
Evergreen tree up to 20 high. Crown at
first pyramidal, then broadly branched
in mature trees. Leaves scale-like, very
small. Fruit a berry-like cone with 5-9
seeds. Native to Balkans, Turkey and
eastward as far as Afghanistan. In-
digenous to Cyprus, restricted to the
Troodos range (1200-1610 m alt.).

TWIG ANATOMY
Bark
Groups of sieve cell in radial rows.
Collapsed sieve cells present. Distinct
ray dilatation in cortex. Sclerenchyma
cells in phloem and in cortex. Fibers in
tangential rows in phloem, scatter and
irregularly dispersed in cortex. Crystal
sand present. With large resin ducts in
cortex. Phellem homogeneous, distinct
in polarized light. Layered phellem.
Junipers excelsa M. Bieb. subsp. excelsa

Xylem
As stem wood.

›T

Pith
Pith shape triangular. Thick-walled
parenchyma cells present. Pits in
transverse and in longitudinal cell
walls. Vascular bundles not distinct.

›T

56
Stem xylem: 40 44 48 53 55 56 72 73 78 80 86 87 93 98 102 107 111
Twig bark: B3 B4 B5 B7 B9 B12 B23 B25 B26 B31 B33 B29
Pith: P1.2 P3 P3.3 P9 P9.1 P10.2

Cupressaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct.
Intercellular spaces throughout the
wood visible in transverse section,
mainly in earlywood. Latewood trac-
heids thick-walled (double wall thick-
ness larger than radial lumen diame-
ter). Axial parenchyma present, diffuse.
Traumatic resin canals present.

›T ›T

Radial section
Pitting in radial walls of earlywood trac-
heids predominantly uniseriate. Organ-
ic deposit in heartwood tracheids pre-
sent. Axial parenchyma transverse end
walls nodular. Ray tracheids absent or
very rare. End walls of ray parenchyma
cells distinctly pitted. Horizontal walls
of ray parenchyma cells smooth (un-
pitted). Cross-field pitting cupressoid.
Number of pits per cross-field in early-
wood 1-3.

Junipers excelsa M. Bieb. subsp. excelsa


›T ›T

Tangential section
Average ray height very low (up to 9
cells). Rays exclusively uniseriate.

Wood basic density: 0,46 g/cm³

›T ›T

57
Juniperus foetidissima Willd.
Stincking Juniper, Foetid Juniper
Cupressaceae

PLANT DESCRIPTION
Evergreen tree up to 20 m high, with
conical or pyramidal crown. Leaves
scale-like, very small, awl-shaped in
young shoots. Fruit a berry-like cone
with 1-3 seeds. Native to Greece, Tur-
key, Caucasia and Syria. Indigenous
to Cyprus, restricted to the Troodos
range (1400-1950 m alt.).

TWIG ANATOMY
Bark
Groups of sieve cell in tangential rows.
Sclerenchyma cells in phloem and in
cortex. Fibers in tangential rows in
phloem, scattered or irregularly dis-
persed in cortex. Crystal sand present.
Phellem homogeneous, distinct in
polarized light. Layered phellem.

Xylem
As stem wood.

›T

Pith
Juniperus foetidissima Willd.

Pith shape round to elongated. Thick-


walled parenchyma cells present. Pits
of two distinct size in transverse and in
longitudinal cell walls. Vascular bun-
dles not clearly distinct.

›T ›T

58
Stem xylem: 40 44 48 54 56 72 73 74 76 80 86 88 93 98 102 103 107
Twig bark: B2 B7 B9 B12 B13 B23 B31 B33 B29 B35
Pith: P1 P3.3 P9 P9.1 P9.3 P10.2

Cupressaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct. Late-
wood tracheids thin-walled (double
wall thickness less than radial lumen
diameter). Axial parenchyma present,
diffuse and tangentially zonate.

›T ›T

Radial section
Pitting in radial walls of earlywood
tracheids predominantly uniseriate.
Organic deposit in heartwood tracheids
present. Axial parenchyma transverse
end walls smooth. Ray tracheids absent
or very rare. End walls of ray parenchy-
ma cells distinctly pitted. Horizontal
walls of ray parenchyma cells distinctly
pitted. Cross-field pitting cupressoid.
Number of pits per cross-field in early-
wood 1-3.

›T ›T

Tangential section Juniperus foetidissima Willd.


Average ray height very low (up to 4
cells) to medium (5 to 15 cells). Rays
exclusively uniseriate.

Wood basic density: 0,69 g/cm³

›T ›T

59
Juniperus oxycedrus L.
Prickly Juniper
Cupressaceae

PLANT DESCRIPTION
Evergreen shrub or tree up to 8 m high,
with rounded or conical crown. Leaves
in whorls of 3, awl-shaped. Fruit a ber-
ry-like cone, reddish-brown. Indigenous
to Cyprus, growing on rocky mountain-
sides (700-1500 m alt.). Widely distrib-
uted from southern Europe eastward to
the Caucasus and Iran.

TWIG ANATOMY
Bark
Groups of sieve cell in radial rows.
Collapsed sieve cells present. Distinct
ray dilatation. Sclerenchyma cells in
phloem and in cortex. Fibers in tan-
gential rows in phloem, scattered and
irregularly dispersed in cortex. Crystal
sand present. Ducts in cortex. Phellem
homogeneous, distinct in polarized
light.

Xylem
As stem wood.

›T

Pith
Pith shape round. Heterogeneous pith.
Thick-walled parenchyma cells present.
Cell content present (dark staining
substances). Pits in transverse and in
Juniperus oxycedrus L.

longitudinal cell walls. Vascular bun-


dles not distinct.

›T

60
Stem xylem: 40 44 48 55 56 72 73 74 79 86 93 98 102 107
Twig bark: B5 B7 B9 B12 B13 B23 B25 B26 B28 B31 B33
Pith: P1 P3 P3.3 P5 P9 P9.1 P10.2

Cupressaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct.
Latewood tracheids thick-walled (dou-
ble wall thickness larger than radial
lumen diameter). Axial parenchyma
present, diffuse and tangentially zon-
ate.

›T ›T

Radial section
Pitting in radial walls of earlywood
tracheids predominantly uniseriate.
Organic deposit in heartwood tracheids
present. Ray tracheids commonly pre-
sent. End walls of ray parenchyma cells
distinctly pitted. Cross-field pitting
cupressoid. Number of pits per cross-
field in earlywood 1-3.

›T ›T

Tangential section
Average ray height very low (up to 4
cells). Rays exclusively uniseriate.
Juniperus oxycedrus L.

Wood basic density: 0,56 g/cm³

›T ›T

61
Juniperus phoenicea L.
Phoenician Juniper
Cupressaceae

PLANT DESCRIPTION
Evergreen shrub or tree up to 10 high,
with conical crown. Leaves scale-like,
very small and awl-shaped in young
shoots. Fruit a berry-like cone, dark
reddish-brown. Widely distributed in
the Mediterranean region. Indigenous
to Cyprus, occurring in coastal maquis
and sand dunes (0-500 m alt.).

TWIG ANATOMY
Bark
Groups of sieve cell and collapsed
sieve cells present. Some rays become
dilated. Sclerenchyma cells in phloem
and in cortex. Fibers present, in tan-
gential lines in phloem, scattered or
irregularly dispersed in cortex. Crystal
sand present. Phellem homogeneous,
distinct in polarized light. Layered
phellem.

Xylem
As stem wood.

›T

Pith
Pith shape polygonal. Thick-walled
parenchyma cells present. Pits in trans-
verse and in longitudinal cell walls.
Vascular bundles not distinct.
Juniperus phoenicea L.

›T

62
Stem xylem: 40 44 55 56 72 74 76 80 86 88 93 98 102 107
Twig bark: B1 B4 B6 B7 B9 B12 B13 B23 B31 B33 B29
Pith: P1.1 P3.3 P9 P9.1 P10.2

Cupressaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct. Late-
wood tracheids thick-walled (double
wall thickness larger than radial lumen
diameter). Axial parenchyma present,
tangentially zonate.

›T ›T

Radial section
Pitting in radial walls of earlywood
tracheids predominantly uniseriate.
Axial parenchyma transverse end walls
smooth. Ray tracheids absent or very
rare. End and horizontal walls of ray
parenchyma cells distinctly pitted.
Cross-field pitting cupressoid. Number
of pits per cross-field in earlywood 1-3.

›T ›T

Tangential section
Average ray height very low (up to 4
cells). Rays exclusively uniseriate.
Juniperus phoenicea L.

Wood basic density: 0,67 g/cm³

›T ›T

63
Cedrus brevifolia (Hook. f.) A. Henry
Cyprus Cedar
Pinaceae

PLANT DESCRIPTION
Evergreen tree up to 30 m high. Crown
at first pyramidal, gradually becoming
broad, with characteristic horizontal
branching in older trees. Leaves needle-
like, whorled, 10-15 mm in length.
Cones woody barrel-shaped. Endemic to
Cyprus, restricted to the Cedar Valley
(900-1400 m alt.).

TWIG ANATOMY
Bark
Collapsed sieve cells present. Scleren-
chyma cells in phloem and in cortex.
Sclereids and fibers scattered or irreg-
ularly dispersed. Prismatic crystals and
crystal sand present. With resin canals
in cortex. Cell content in parenchyma
cells. Phellem distinct in polarized
light.

Xylem
As stem wood.
Cedrus brevifolia (Hook. f.) Henry

Pith
Pith shape polygonal. Heterogeneous
pith. Thick-walled parenchyma cells
present. Unlignified cells present. Cell
content present (dark staining sub-
stances). Prismatic crystals present.
Pits in transverse and in longitudinal
cell walls. Vascular bundles not dis-
tinct.

64
Stem xylem: 40 44 55 57 79 81 86 88 94 98 104 107
Twig bark: B4 B7 B15 B18 B20 B23 B25 B26 B28 B33
Pith: P1.1 P2 P3 P3.3 P3.4 P5 P6 P9 P9.1 P10.2

Pinaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct. Late-
wood tracheids thick-walled (double
wall thickness larger than radial lumen
diameter).

Radial section
Pitting in radial walls of earlywood tra-
cheids predominantly uniseriate. Torus
in earlywood tracheids’ pits scalloped.
Ray tracheids commonly present. Cell
walls of ray tracheids smooth. End
walls of ray parenchyma cells distinctly
pitted. Horizontal walls of ray paren-
chyma cells distinctly pitted. Cross-
field pitting taxodioid. Number of pits
per cross-field in earlywood 1-3.

Tangential section Cedrus brevifolia (Hook. f.) Henry


Average ray height high (from 16 to 30
cells). Rays exclusively uniseriate.

Wood basic density: 0,51 g/cm³

65
Cedrus libani A.Rich.
Lebanese Cedar
Pinaceae

PLANT DESCRIPTION
An evergreen tree growing up to 40 m
high. The crown is conic when young,
becoming broadly tabular with age
with fairly level branches. The leaves
are needle-like, spaced out on the long
shoots, and in clusters of 15-45 on the
shoots; they are 5-30 mm in length. On
Cyprus it grows at 1000-1525 m alt.

Photo: Ori Fragman-Sapir


TWIG ANATOMY
Bark
Groups of sieve cell in tangential rows.
Sclerenchyma cells in phloem and in
cortex. Fibers in tangential rows in
phloem. Sclereids scattered or irregu-
larly dispersed in cortex. Prismatic
crystals present. Phellem homogene-
ous, distinct in polarized light. Ligni-
fied cells in phellem.

Xylem
As stem wood.

›T

Pith
Pith shape polygonal. Thick-walled
parenchyma cells present. Cells dimor-
phic. Pits in transverse and in longi-
tudinal cell walls. Vascular bundles
clearly separate. Axial cells in regular
Cedrus libani A.Rich.

rows (radial section).

›T

66
Stem xylem: 40 44 50 55 56 57 79 81 86 88 94 98 104 105 107 108 111 118 119 122
Twig bark: B2 B7 B9 B12 B15 B18 B20 B31 B33 B34
Pith: P1.1 P3.3 P4 P9 P9.1 P10.1 P13

Pinaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct. Late-
wood tracheids thick-walled (double
wall thickness larger than radial lu-
men diameter). Traumatic resin canals
present.

›T ›T

Radial section
Pitting in radial walls of earlywood
tracheids predominantly uniseriate.
Average tracheids length short (less
than 3000 μm). Torus in earlywood
tracheids’ pits scalloped. Ray trac-
heids commonly present. Cell walls
of ray tracheids smooth. End walls of
ray parenchyma cells distinctly pitted.
Horizontal walls of ray parenchyma
cells distinctly pitted. Cross-field pit-
ting taxodioid, number of pits per
cross-field in earlywood 1-3. Prismatic
crystals in rays.

›T ›T

Tangential section
Average ray height high to very high
(more than 30 cells). Rays uniseriate
and in part 2-3 seriate.
Cedrus libani A.Rich.

Wood basic density: 0,45 g/cm³

›T ›T

67
Pinus brutia Ten.
Calabrian Pine
Pinaceae

PLANT DESCRIPTION
Evergreen, resinous tree up to 40 m
high. Leaves needle-like, bright green
or greyish–green, 11-15 cm long. In-
digenous to Cyprus, it is the dominant
forest species on the island, with a
wide distribution on a variety of habi-
tats (0-1400 m alt.). Native to southern
Italy, eastward to northern Iran.

TWIG ANATOMY
Bark
Groups of sieve cell and collapsed
sieve cells present. Sclerenchyma cells
in phloem and in cortex. Prismatic
crystals present. With resin ducts cor-
tex. Cell content in parenchyma cells.
Epidermis distinct in polarized light.

Xylem
As stem wood.

›T

Pith
Pith shape polygonal. Heterogenous
pith, Sclereids present. Prismatic crys-
tals present. Pits in transverse and in
longitudinal cell walls. Vascular bun-
dles not distinct.
Pinus brutia Ten.

›T ›T

68
Stem xylem: 40 44 54 56 79 82 86 88 93 94 98 103 107 109 110 117 118 119 124
Twig bark: B1 B4 B7 B20 B25 B26 B28 B31 B33
Pith: P1.1 P3 P3.1 P6 P9 P9.1 P10.2

Pinaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct. Late-
wood tracheids thin-walled (double wall
thickness less than radial lumen diam-
eter). Axial intercellular resin canals
present. Epithelial cells thin-walled.

›T ›T

Radial section
Pitting in radial walls of earlywood tra-
cheids predominantly uniseriate. Ray
tracheids commonly present. Cell walls
of ray tracheids dentate. End walls of
ray parenchyma cells distinctly pitted.
Horizontal walls of ray parenchyma
cells distinctly pitted. Cross-field pit-
ting cupressoid to taxodioid. Number
of pits per cross-field in earlywood 1-3.
Prismatic crystals located in cells asso-
ciated with intercellular canals.

›T ›T

Tangential section
Average ray height medium (5 to 15
cells). Rays exclusively uniseriate. Ra-
dial intercellular resin canals present.
Pinus brutia Ten.

Wood basic density: 0,46 g/cm³

›T ›T

69
Pinus halepensis L.
Aleppo Pine, Jerusalem Pine
Pinaceae

PLANT DESCRIPTION
A small to medium-size tree, 15–25 m
high. The needle-like leaves are very
slender, distinctly yellowish green and
produced in pairs. The cones are nar-
row conic, 5–12 cm long. It generally
grows at low altitudes, mostly from sea
level to 200 meters, but can grow at an
altitude of up to 1000 m.

TWIG ANATOMY
Bark
Groups of sieve cell in tangential rows.
Sclerenchyma cells in phloem and in
cortex. Fibers scattered or irregularly
dispersed. Prismatic crystals present.
With ducts in cortex.

Xylem
As stem wood.

Pith
No slide available.
Pinus halepensis L.

›T

70
Stem xylem: 40 44 55 56 79 82 86 88 91 98 103 107 109 110 117
Twig bark: B1 B2 B7 B9 B13 B20 B25 B26 B31 B33
Pith: No slide available.

Pinaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct. Late-
wood tracheids thick-walled (double
wall thickness larger than radial lu-
men diameter). Axial intercellular resin
canals present. Epithelial cells thin-
walled.

›T ›T

Radial section
Pitting in radial walls of earlywood tra-
cheids predominantly uniseriate. Ray
tracheids commonly present. Cell walls
of ray tracheids dentate. End walls of
ray parenchyma cells distinctly pitted.
Horizontal walls of ray parenchyma
cells distinctly pitted. Cross-field pit-
ting pinoid. Number of pits per cross-
field in earlywood 1-3.

›T ›T

Tangential section
Average ray height medium (5 to 15
cells). Rays exclusively uniseriate. Ra-
dial intercellular resin canals present.
Pinus halepensis L.

Wood basic density: 0,46 g/cm³

›T ›T

71
Pinus nigra subsp. pallasiana (D.Don) Holmboe
Black Pine, Palla’s Pine
Pinaceae

PLANT DESCRIPTION
Evergreen tree up to 40 m high. Leaves
spirally arranged, of two kinds: scale-
like on long shoots and needle-like,
paired, in short shoots. Indigenous
to Cyprus, occurring on the highest
slopes of Troodos (1100-1950 m alt.).
It occurs in the Balkan Peninsula, Tur-
key, Crimea and the Carpathians.

TWIG ANATOMY
Bark
Groups of sieve cell in tangential rows.
Collapsed sieve cells present. Scleren-
chyma cells in phloem and in cortex.
Prismatic crystals present. With resin
ducts in cortex. Cell content in paren-
Pinus nigra subsp. pallasiana (D.Don) Holmboe

chyma cells. Phellem homogeneous,


distinct in polarized light.

Xylem
As stem wood.

›T

Pith
Pith shape polygonal. Heterogenous
pith. Sclereids present. Pits in trans-
verse and in longitudinal cell walls.
Vascular bundles clearly separate.

›T ›T

72
Stem xylem: 40 44 55 56 79 82 85 87 90 97 103 107 109 110 117 118 119 124
Twig bark: B2 B4 B7 B20 B25 B26 B28 B31 B33
Pith: P1.1 P3 P3.1 P9 P9.1 P10.1

Pinaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct. Late-
wood tracheids thick-walled (double
wall thickness larger than radial lumen
diameter). Axial intercellular resin ca-
nals present with epithelial cells thin-
walled.

›T ›T

Radial section
Pitting in radial walls of earlywood tra-
cheids predominantly uniseriate. Ray
tracheids commonly present. Cell walls
of ray tracheids dentate. End walls of
ray parenchyma cells smooth. Hori-
zontal walls of ray parenchyma cells
smooth (unpitted). Cross-field pitting

Pinus nigra subsp. pallasiana (D.Don) Holmboe


fenestriform (“window-like”). Number
of pits per cross-field in earlywood 1-2
(large fenestriform). Prismatic crystals
located in cells associated with inter-
cellular canals.

›T ›T

Tangential section
Average ray height medium (5 to 15
cells). Rays exclusively uniseriate. Ra-
dial intercellular resin canals present.

Wood basic density: 0,45 g/cm³

›T ›T

73
Pinus pinea L.
Umbrella Pine
Pinaceae

PLANT DESCRIPTION
Evergreen tree up to 30 m high, with a
characteristic spherical crown. Leaves
needle-like, 8-12 cm long, found in
pairs, bright green. Cone woody, large,
broadly ovate. Exotic to Cyprus, widely
cultivated in gardens and plantations
(0-140 m alt.). Native of the Mediterra-
nean region and Portugal.

TWIG ANATOMY
Bark
Sclerenchyma cells in phloem and in
cortex. Prismatic crystals present. Cor-
tex consist of large and small paren-
chyma cells. With resin ducts in cor-
tex. Cell content in parenchyma cells.
Epidermis distinct in polarized light.

Xylem
As stem wood.

›T

Pith
Pith shape polygonal. Cell content
present (dark staining substances).
Prismatic crystals present. Pits in trans-
verse and in longitudinal cell walls.
Vascular bundles not clearly distinct.
Resin duct at the periphery of the pith.
Pinus pinea L.

›T ›T

74
Stem xylem: 40 44 54 56 79 81 86 88 93 98 103 107 109 110 117
Twig bark: B7 B20 B25 B26 B28 B31 B33
Pith: P1.1 P5 P6 P9 P9.1 P10.2

Pinaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct.
Latewood tracheids thin-walled (double
wall thickness less than radial lumen
diameter). Axial intercellular resin
canals present. Epithelial cells thin-
walled.

›T ›T

Radial section
Pitting in radial walls of earlywood tra-
cheids predominantly uniseriate. Ray
tracheids commonly present. Cell walls
of ray tracheids smooth. End walls of
ray parenchyma cells distinctly pitted.
Horizontal walls of ray parenchyma
cells distinctly pitted. Cross-field pit-
ting cupressoid. Number of pits per
cross-field in earlywood 1-3.

›T ›T

Tangential section
Average ray height medium (5 to 15
cells). Rays exclusively uniseriate. Ra-
dial intercellular resin canals present.
Pinus pinea L.

Wood basic density: 0,45 g/cm³

›T ›T

75
Taxus baccata L.
Common Yew, English Yew
Taxaceae

PLANT DESCRIPTION
Evergreen tree up to 15 m high, with
broad crown and reddish-brown bark.
Leaves linear, flat, 2-3 cm long. An
exotic species to Cyprus, cultivated as
ornamental in gardens and parks (500-
1000 m alt.). Native to Europe, north
Africa and western Asia, only on moun-
tains in the Mediterranean region.

TWIG ANATOMY
Bark
Groups of sieve cell in tangential rows.
Sclerenchyma cells in phloem and in
cortex. Fibers scattered or irregularly
dispersed. Crystal sand present. Phel-
lem homogeneous, distinct in polar-
ized light. Lignified cells in phellem.

Xylem
As stem wood.

›T

Pith
Pith not visible in polarized light. Pith
shape polygonal. Unlignified cells
present. Cells dimorphic. Crystal sand
present. Vascular bundles clearly sepa-
rate.
Taxus baccata L.

›T

76
Stem xylem: 40 44 55 56 61 62 65 68 80 85 87 93 98 103 107
Twig bark: B1 B2 B7 B9 B13 B23 B31 B33 B34
Pith: P0.1 P1.1 P3.4 P4 P6.4 P10.1

Taxaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct. Late-
wood tracheids thick-walled (double
wall thickness larger than radial lumen
diameter).

›T

Radial section
Pitting in radial walls of earlywood tra-
cheids predominantly uniseriate. Heli-
cal thickenings in tracheids present.
Helical thickenings single and widely
spaced (number of coils less than 120
per mm). Ray tracheids absent or very
rare. End walls of ray parenchyma cells
smooth. Horizontal walls of ray paren-
chyma cells smooth (unpitted). Cross-
field pitting cupressoid. Number of pits
per cross-field in earlywood 1-3.

›T ›T

Tangential section
Average ray height medium (5 to 15
cells). Rays exclusively uniseriate.
Taxus baccata L.

Wood basic density: 0,52 g/cm³

›T ›T

77
Gnetales

This section contains anatomical


descriptions of stem and twig xylem,
as well as the bark and pith regions of
two gnetales species belonging to the
Ephedraceae family.

Gnetales
Ephedra foeminea Forssk. = Ephedra fragilis Desf subsp.
campylopoda (C.A. Mey.) Asch. et Graebn.
Ephedraceae

PLANT DESCRIPTION
Dioecious, erect or climbing, much
branched shrub, with a stem up to 3 m
long. Branches pendulous, glaucous-
green. Native species in the eastern
Mediterranean basin. Fairly common in
Cyprus, occurring in maquis, garigue,
stone walls and on rocky slopes (0-900
m alt.).

TWIG ANATOMY
Bark
Groups of sieve cell and collapsed
sieve cells present. Only some rays be-
come slightly dilated. Cortex consist of
large thin-walled parenchyma cells and
scattered or irregularly dispersed fib-
ers and sclereids. Crystal sand present.
Epidermis distinct in polarized light.

Xylem
Thick-walled fibers in the first ring.
Vessel clusters common. Intervessel
pits minute (less than 4 μm in diam-
eter). Rays uniseriate.
›T

Pith
Pith shape roundish. Large cells in the
centre, smaller at the border of the
Ephedra foeminea Forssk.

pith. Cell content present (dark stain-


ing substances). Vascular bundles
clearly separate.

›T ›T

80
Stem xylem: 1 4 9 19 26 30 40.2 50.2 53.2 58 62 69 70.3 75 78 97 105 109 115
Twig bark: B1 B4 B6 B9 B13 B15 B18 B23 B31 B33
Pith: P1.1 P4.1 P5 P10.1

Ephedraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood semi-ring-porous, vessels
predominantly solitary. Mean tangen-
tial diameter of earlywood vessels
20-50 μm, more than 200 vessels per
mm2. Fibers thin- to thick-walled, radial
flat marginal fibers. Axial parenchyma
extremely rare or not to recognizable,
sometimes scanty paratracheal. Rays
4-12 per mm.

›T ›T

Radial section
Foraminate perforation plates. Interves-
sel pits medium (7-10 μm in diameter).
Vessel-ray pits half bordered, smaller
than intervessel pits. Earlywood ves-
sel element length more than 500 μm.
Fibers with distinctly bordered pits
(fiber tracheids). All ray cells upright or
square, with some procumbent cells

›T ›T

Tangential section
Ray width predominantly 1 to 3 cells.
Dark staining substances in ray cells.
Ephedra foeminea Forssk.

Wood basic density: 0,73 g/cm³

›T ›T

81
Ephedra nebrodensis Tineo subsp. procera (Fisch.
et C.A. Mey.) K. Richt.
Ephedraceae

PLANT DESCRIPTION
Dioecious, erect, scoparioid shrub up
to 1 m high. Branches erect, glaucous-
green. Indigenous to the Mediterrane-
an countries and the countries east-
ward to Iran, Afghanistan and central
Asia. Rare in Cyprus, it occurs on rocky
slopes, in maquis and pine forest
openings (600-900 m alt.).
Ephedra nebrodensis Tineo subsp. procera (Fisch. et C.A. Mey.) K. Richt.

Photo: Hadjikyriakou
TWIG ANATOMY
Bark
Groups of sieve cell and collapsed
sieve cells present. Sclerenchyma cells
in phloem and in cortex. Sclereids scat-
tered or irregularly dispersed. Crystal
sand present. Cell content in parenchy-
ma cells. Layered phelloderm.

Xylem
Rays per mm 12-20.

›T

Pith
Pith shape round. Medullary sheath
present. Cells dimorphic. Cell content
present (dark staining substances).
Vascular bundles clearly separate from
one other to not distinct. Axial cells in
regular rows (radial section).

›T ›T

82
Stem xylem: 1 3 9 19 27 30 40.1 47 53.2 62 69 76 78 86 98 105 115 152
Twig bark: B1 B4 B7 B18 B23 B28 B35
Pith: P1 P2 P4 P5 P10.1 P10.2 P13

Ephedraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood ring-porous. Vessels
predominantly solitary. Mean tangen-
tial diameter of earlywood vessels less
than 20 μm, 5-20 vessels per mm2. Fib-
ers thin- to thick-walled. Axial paren-
chyma diffuse and scanty paratracheal.
Apotracheal parenchyma in narrow

Ephedra nebrodensis Tineo subsp. procera (Fisch. et C.A. Mey.) K. Richt.


bands or lines up to three cells wide.
Rays 4-12 per mm.

›T ›T

Radial section
Foraminate perforation plates. Interves-
sel pits large (more than 10 μm in
diameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length more
than 500 μm. Fibers with distinctly bor-
dered pits (fiber tracheids). All ray cells
upright and square. Crystal sand.

›T ›T

Tangential section
Ray width predominantly 1 to 5 cells.

Wood basic density: 0,83 g/cm³

›T ›T

83
Angiosperms: Monocotyledons

This section contains anatomical


descriptions of stem and bark of 6
monocotyledonous species belonging
to 4 families.

Angiosperms: Monocotyledons
Asparagus acutifolius L.
Hardy Asparagus
Asparagaceae

PLANT DESCRIPTION
Monocot with angular, often purplish
stems up to 2 m long. Leaves alternate
membranous, scaly. Cladodes in fasci-
cles of 5-30, spiny. A common indig-
enous species to Cyprus, occurring on
maquis, garigue and pine forests (0-
1500 m alt.). Indigenous to European
and Mediterranean countries.

Photo: Hadjikyriakou
STEM ANATOMY
Circular outline. Thin-walled unligni-
fied epidermis with a thin cuticle. Be-
low a 4-6 cell wide thin-walled, unligni-
fied assimilatory tissue is a lignified
sclerenchymatic belt, which changes
continuously or abruptly into a thin–to
thick-walled tissue up to the filled cen-
tre. Horseshoe-like vascular bundles
are accompanied by a one-sided small
layer of Unlignified cells present. Two
large vessels with a diameter of 80 to
100 μm and a group of horseshoe like
arranged small vessels are arranged
around the phloem.
›T
Asparagus acutifolius L.

Stem basic density: 0,60 g/cm³

›T ›T

86
Asparagus stipularis Forsskal
Wild Asparagus

Asparagaceae
PLANT DESCRIPTION
Erect or sprawling shrub up to 2 m
long. Stem angular, glaucous-green.
Laves alternate, scaly, membranous.
Common, indigenous species to Cy-
prus that occur in maquis, garigue and
pine forests (0-600 m alt.). Indigenous
also to Mediterranean countries and
Atlantic islands.

STEM ANATOMY
Round to slightly wavy outline. Thin-
walled unlignified epidermis with a
thick cuticle. Below a 3-5 cell wide
thin-walled, unlignified assimilatory
tissue is a lignified sclerenchymatic
belt, which changes fairly abrupt into
a thin- to thick-walled tissue up to the
filled centre. Heart-like thin-walled
vascular bundles are accompanied by
a one-sided small layer of Unlignified
cells present. Two large vessels with a
diameter of 80 to 100 μm and a group
of horseshoe like arranged small ves-
sels are arranged around the phloem.
›T

Asparagus stipularis Forsskal

Stem basic density: 0,58 g/cm³

›T ›T

87
Arundo donax L.
Giant Reed
Graminaceae

PLANT DESCRIPTION
Perennial herb up to 6 m high with
stout, long and woody rhizome. Culms
woody, erect, hollow, 2-4 cm in diam-
eter. Leaves linear up to 60x8 cm. It
occurs in moist places along rivers,
streams and canals (0-1400 m alt.). It
also occurs in the Atlantic islands, tropi-
cal Asia eastward to New Zealand.

STEM ANATOMY
Outline circular. The periphery consists
on thin cuticula, very small epidermis
cells and a thick-walled hypodermis.
Below is a 5-8 cell wide homogenous,
thick-walled lignified assimilatory
tissue and an extremely thick-walled
lignified sclerenchymatic belt, which
changes fairly abrupt into a thin- to
thick-walled tissue up to hollow centre.
Round to heart-like vascular bundles
are surrounded by a large layer of
thick-walled lignified cells. Two large
vessels with a diameter of 80 to 100
μm are placed lateral of a round phlo-
em. Characteristic is a radial row of ›T
thick-walled vessels of the protoxylem
between the lateral vessels.
Arundo donax L.

Stem basic density: 0,62 g/cm³

›T

88
Pharagmites australis (Cav.) Trin. ex Steud
Common Reed

Graminaceae
PLANT DESCRIPTION
Perennial herb up to 3 m high, spread-
ing principally by rhizome. Culms
woody, erect, hollow, 1-2 cm in di-
ameter. Leaves linear up to 40x4 cm.
Indigenous to Cyprus, found in moist
places, along rivers, streams and saline
marshes (0-600 m alt.). It also occurs in
many temperate and tropical countries.

STEM ANATOMY
Outline circular. Very thick-walled un-
lignified epidermis consists on quad-
rangular cells which are covered with
a thin cuticula. Below is a two-layered
thick-walled hypodermis and a thin- to
thick-walled assimilatory tissue. The
very thick-walled sclerenchymatic belt
changes fairly abrupt towards a fairly
thin-walled tissue up to large hollow

Pharagmites australis (Cav.) Trin. ex Steud


centre. Tangential oval vascular bun-
dles are surrounded by a thin layer of
lignified thick-walled cells. Two large
vessels with a diameter of 100 to
130 μm are placed lateral of a round
›T phloem. Typical is a thick-walled vessel
of the protoxylem between the lateral
vessels.

Stem basic density: 0,73 g/cm³

›T
›T

89
Ruscus aculeatus L.
Butcher’s Broom
Ruscaceae

PLANT DESCRIPTION
Evergreen shrub with stems up to 80
cm long. Rhizome compact and short.
Stem dark green, glabrous. Leaves
alternate, scaly, membranous. Rather
uncommon in Cyprus, found in hedges
and shrubland (0-1100 m alt.). It also
occurs in Europe, and in the Mediterra-
nean countries.

STEM ANATOMY
Outline circular with rips. Thin-walled
unlignified epidermis with a thick, fine
ripped cuticula. Below 10-15 cell wide
heterogenous, unlignified assimilatory
tissue is a lignified sclerenchymatic
belt, which changes fairly abrupt into a
fairly thin-walled tissue up to the filled
centre. The assimilatory belt is delimi-
tated toward the sclerenchymatic belt
by an endodermis. Round thin-walled
vascular bundles are accompanied
at the peripheral side by a group of
half-moon-like thick-walled fibers and
a row of thin- to thick-walled fibers at
the centripetal side. Groups of small
›T
vessels with a diameter of 10 to 20 μm
are placed half-moon-like at the cen-
tripetal side of a round to oval phloem.
Ruscus aculeatus L.

Stem basic density: 0,52 g/cm³

›T ›T

90
Smilax aspera L.
Common Smilax

Smilaceae
PLANT DESCRIPTION
Evergreen, climbing shrub with stems
up to 4 m long, with dense, patent or
recurved prickles. Leaves cordate and
leathery. Flowers small, yellowish; fruit
a red, globose berry. A Mediterranean
species, in Cyprus it grows in maquis,
garigue and pine forests (0-1400 m
alt.).

STEM ANATOMY
Round to slightly wavy outline. Thin-
walled unlignified epidermis with a
thick cuticula. Below a 7-10 cell wide
thin- to thick-walled, lignified assimila-
tory tissue is a lignified sclerenchym-
atic belt, which changes fairly abrupt
into a fairly thin-walled tissue up to the
filled centre. Round vascular bundles
are accompanied by a layer of lignifed
thin- to thick-walled cells. Two large
vessels with a diameter of 100 to 150
μm and a group of few small vessels
are arranged centripetal below a round
phloem.
›T

Smilax aspera L.

Stem basic density: 0,67 g/cm³

›T

91
Angiosperms: Dicotyledons

This section contains anatomical de-


scriptions of stem and twig xylem, as
well as the bark and pith regions of
244 dicotyledonous species belonging
to 61 families.

Angiosperms: Dicotyledons
Sambucus nigra L.
Common Elder, Black Elder
Adoxaceae

PLANT DESCRIPTION
Deciduous shrub up to 10 m high.
Bark grey-brown. Leaves opposite,
compound, imparipinnate. Natural-
ized in Cyprus, found in field margins,
river banks and home gardens (0-1200
m alt.). It also occurs throughout the
Mediterranean Basin, in central and
northern Europe and western Asia.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclerenchyma cells
in phloem and in cortex. Fibers scat-
tered or irregularly dispersed. Crystal
sand present. Cortex homogeneous.
Epidermis distinct in polarized light.

Xylem
Vessel solitary and in small clusters.
Mean tangential diameter of earlywood
vessel lumina 20-50 μm. Rays predom-
inantly uniseriate.

›T

Pith
Pith shape round. Heterogenous pith.
Few scattered thick-walled parenchyma
cells present. Crystal sand present. Pits
in transverse and in longitudinal cell
walls. Vascular bundles clearly sepa-
rate.
Sambucus nigra L.

›T ›T

94
Stem xylem: 1 5 6 11 13 22 25 30 41 50.2 52.3 56 60 61 69 70.2 75 76 97 107 115
Twig bark: B1 B4 B7 B9 B13 B23 B31 B33
Pith: P1 P3 P3.3 P6.4 P9 P9.1 P10.1

Adoxaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted latewood fibers. Wood
diffuse-porous. Vessels predominantly
in clusters, arranged in intra-annual
tangential bands. Mean tangential
diameter of earlywood vessel lumina
50-100 μm. More than 200 vessels per
mm2. Tyloses with thin walls common.
Fibers thin- to thick-walled. Tension
wood present. Axial parenchyma ex-
tremely rare, diffuse. Rays 4-12 per
mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessels length 100-200 μm.
Vascular and/or vasicentric tracheids
present in latewood. Fibers with simple
to minutely bordered pits (libriform
fibers). Body ray cells procumbent with
mostly 2-4 rows of upright and square
marginal cells.

›T ›T

Tangential section
Ray width predominantly 1 to 3 cells.
Sambucus nigra L.

Wood basic density: 0,50 g/cm³

›T ›T

95
Viburnum opulus L.
Guelder Rose, Snowball
Adoxaceae

PLANT DESCRIPTION
Deciduous shrub up to 4 m high.
Leaves opposite, simple, thinly hairy
below. Flowers white in umbellate fas-
cicles. Fruit a sub globose, red drupe.
Exotic to Cyprus, found in gardens and
hedges and elsewhere (500-1200 m
alt.). Indigenous to central and south-
ern Europe.

TWIG ANATOMY
Bark
Groups of sieve tube present. Scleren-
chyma cells in phloem and in cortex.
Sclereids scattered or irregularly dis-
persed and in small groups. Crystal
druses present. Cortex and phellem
homogeneous, the latter distinct in
polarized light. Layered phelloderm.

Xylem
As stem wood.

›T

Pith
Pith shape hexagonal. Medullary
sheath present. Cells dimorphic. Crys-
tal druses present. Pits in transverse
and in longitudinal cell walls, grouped.
Pits of two distinct size. Vascular bun-
dles clearly separate to not distinct.
Viburnum opulus L.

Tracheary elements of metaxylem in


distinct radial rows. Axial cells in regu-
lar rows (radial section).

›T ›T

96
Stem xylem: 1 5 9 11 14 18 21 25 30 40.2 50.2 53.2 62 69 70.2 76 78 96 109 116.2
Twig bark: B1 B4 B7 B18 B19 B22 B31 B33 B35
Pith: P1.1 P2 P4 P6.2 P9 P9.1 P9.2 P9.3 P10.1 P10.2 P12 P13

Adoxaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood diffuse-porous. Vessel
solitary or in small clusters. Mean tan-
gential diameter of earlywood vessels
20-50 μm. More than 200 vessels per
mm2. Fibers thin- to thick-walled. Ten-
sion wood present. Axial parenchyma
diffuse and scanty paratracheal. More
than 20 rays per mm.

›T ›T

Radial section
Scalariform perforation plates with
more than 40 bars. Inter-vessel pits
opposite, small (4-7 μm in diameter).
Vessel-ray pits with distinct borders,
similar to intervessel pits in size and
shape throughout the ray cell. Early-
wood vessel element length more than
500 μm. Fibers with distinctly bor-
dered pits (fiber tracheids). Rays with
procumbent, square and upright cells
mixed throughout the ray.

›T ›T

Tangential section
Rays exclusively uniseriate.
Viburnum opulus L.

Wood basic density: 0,50 g/cm³

›T ›T

97
Viburnum tinus L. subsp. tinus
Laurustinus
Adoxaceae

PLANT DESCRIPTION
Evergreen shrub up to 7 m high. Leaves
opposite, leathery, thinly hairy below.
Flowers white in umbellate fascicles.
Fruit a sub-globose, red drupe. A rare
indigenous to Cyprus recently found on
Pentadaktylos range; also planted in gar-
dens and hedges and elsewhere (0-1400
m alt.). Indigenous to southern Europe.

TWIG ANATOMY
Bark
Groups of sieve tube present. Fibers
scattered or irregularly dispersed in
cortex. Prismatic crystals and crystal
druses present. Phellem homogene-
ous. Lignified cells in phellem.

Xylem
Rays exclusively uniseriate.

›T
Viburnum tinus L. subsp. tinus

Pith
Pith shape round. Heterogenous pith.
Thick-walled parenchyma cells present.
Crystal druses present. Pits in trans-
verse and in longitudinal cell walls.
Pits grouped. Vascular bundles clearly
separate to not distinct. Tracheary ele-
ments of metaxylem in distinct radial
rows. Axial cells in regular rows (radial
section).

›T ›T

98
Stem xylem: 1 4 5 9 11 14 17 21 26 30 36 40.2 50.2 53.2 62 70 70.3 76 78 86 96 97 109 116.2
Twig bark: B1 B13 B20 B22 B31 B34
Pith: P1 P3 P3.3 P6.2 P9 P9.1 P9.2 P10.1 P10.2 P12 P13

Adoxaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by a
lack of vessels in terminal latewood
and by distinct by radially flatted and
thick-walled latewood fibers. Wood
diffuse-porous to semi-ring-porous.
Vessels solitary or in small clusters.
Mean tangential diameter of earlywood
vessels 20-50 μm. More than 200 ves-
sels per mm2. Vessel-ray pits with dis-
tinct borders, similar to intervessel pits
in size and shape throughout the ray
cell. Fibers very thick-walled, marginal
flat fibers. Apotracheal parenchyma
diffuse, and in narrow bands or lines
›T ›T up to three cells wide. More than 20
rays per mm.
Radial section
Scalariform perforation plates with
20-40 bars. Intervessel pits opposite,
medium (7-10 μm in diameter). Helical
thickenings in vessel element present.
Earlywood vessel element length more
than 500 μm. Fibers with distinctly bor-
dered pits (fiber tracheids). Rays with
procumbent, square and upright cells
mixed throughout the ray.

›T ›T

Tangential section Viburnum tinus L. subsp. tinus


Rays uniseriate and up to 3 cells wide.

Wood basic density: 0,67 g/cm³

›T ›T

99
Bosea cypria Autran et Schinz
Cyprus Bosea
Amaranthaceae

PLANT DESCRIPTION
Evergreen shrub, 1-2 m high. Fruit
a red globose berry. Leaves broadly
lanceolate to elliptic. Endemic to Cy-
prus, occurring on rocky ground, old
stone walls and rocky sites with open or
maquis vegetation (0-650 m alt.). One
of the three Bosea species occurring
worldwide.

TWIG ANATOMY
Bark
Groups of sieve tube present. Fibers
scattered or irregularly dispersed and
in tangential groups in cortex. Cortex
consist of large and small thin-walled
parenchyma cells. Crystal druses and
crystal sand present. Epidermis distinct
in polarized light.

Xylem
As stem wood.

›T
Bosea cypria Autran et Schinz

Pith
Pith shape round. With prismatic crys-
tals and crystal druses. Pits of two
distinct sizes, grouped in both lon-
gitudinal and transverse cell walls.
Vascular bundles clearly separate to
not distinct. Axial cells in regular rows
(radial section).

›T ›T

100
Stem xylem: 1 5 9 9.1 13 20 22 25 30 40.2 45 50.2 53.1 56 58 61 69 78 89 96 98 102 103 105 116.1 133.1 136 144
Twig bark: B1 B9 B13 B14 B22 B23 B33
Pith: P1 P6 P6.2 P9 P9.1 P9.2 P9.3 P10.1 P10.2 P13

Amaranthaceae
STEM ANATOMY
Transverse section
Growth ring boundaries present in
xylem, distinct by radially flatted and
thick-walled latewood fibers. Wood
diffuse-porous. Vessels solitary or in
short radial multiples. Mean tangential
diameter of earlywood vessels 20-50
μm. More than 200 vessels per mm2.
Vessels of two distinct diameter class-
es.Tyloses with thin walls common.
Deposits in heartwood vessels. Fib-
ers thin- to thick-walled. Parenchyma
scanty paratracheal. Apotracheal pa-
renchyma in marginal bands. Rays 12-
20 per mm. Included phloem: concen-
›T ›T
tric arranged single vascular bundles.
Crystal druses in included phloem.
Radial section
Simple perforation plates. Intervessel
pits pseudoscalariform to reticulate
and alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 200-
500 μm. Fibers with simple to minutely
bordered pits (libriform fibers). Most of
ray cells upright, few squared. Prismat-
ic crystals and crystal druses present.

›T ›T

Tangential section Bosea cypria Autran et Schinz


Rays of two distinct sizes: uniseriate
and 3 to 7 seriate. Wider rays height
more than 1 mm.

Wood basic density: 0,50 g/cm³

›T ›T

101
Cotinus coggygria Scop.
Wig Tree
Anacardiaceae

PLANT DESCRIPTION
Deciduous shrub up to 4 m high,
with resinous odour. Branches glossy-
green at the first, then reddish-brown.
Leaves simple, alternate, oval-elliptical
and greenish-yellow flowers. It grows
throughout the northern Mediterra-
nean countries. Widely cultivated in
gardens (0-900 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Fibers in tangen-
tial bands. Prismatic crystals present.
Secretory elements in ducts in phloem
and in cortex. Phellem homogeneous. ›T

Lignified cells in phellem. Layered


phellem.

Xylem
Mean tangential diameter of earlywood
vessel lumina 20-50 μm. Rays exclu-
sively uniseriate.

›T

Pith
Pith shape round. Cells dimorphic.
Vascular bundles clearly separate.
Tracheary elements of metaxylem in
Cotinus coggygria Scop.

distinct radial rows.

›T ›T

102
Stem xylem: 1 3 11 13 22 25 31 32 36 41 45 49 53.1 56 60 61 69 78 97 106 107 108 115 136 136.1
Twig bark: B1 B4 B12 B20 B25 B26 B31 B34 B29
Pith: P1 P4 P10.1 P12

Anacardiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
the difference in vessel size between
latewood and earlywood. Wood ring-
porous. Vessel clusters common. Mean
tangential diameter of earlywood ves-
sel lumina 50-100 μm, 40-100 vessels
per mm2. Vessel of two distinct diam-
eter classes. Tyloses with thin walls
common. Fibers thin- to thick-walled.
Axial parenchyma scanty paratracheal.
Rays 4-12 per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits rounded or
elongated with large apertures. Helical
thickenings in vessel elements present.
Earlywood vessels length 200-500 μm.
Vascular and/or vasicentric tracheids
present. Fibers with simple to minutely
bordered pits (libriform fibers). Body
ray cells procumbent with one to more
than 4 rows of square marginal cells.
Prismatic crystals present in ray paren-
chyma cells.

›T ›T

Tangential section
Ray width predominantly 1 to 3 cells.
Cotinus coggygria Scop.

Wood basic density: 0,54 g/cm³

›T ›T

103
Pistacia atlantica Desf.
Terebinth Tree
Anacardiaceae

PLANT DESCRIPTION
A robust, deciduous tree up to 15
m high with a massive trunk. Leaves
compound, leaflets in 3-5 pairs, lan-
ceolate or ovate. Fruit an ovate drupe,
at first red, bluish-green at maturity.
Indigenous to Cyprus, in abandoned
fields, fields margins and rocky slopes
(0-1500 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclerenchyma
cells in phloem and in cortex. Fibers
and sclereids in tangential rows and
in groups. Prismatic crystals present.
With secretory elements in ducts. Phel-
lem homogeneous, distinct in polar-
ized light.

Xylem
Vessels in radial multiples of 4 or
more. Vessel diameter less than 20
μm, in two distinct diameter classes.
Some ray with procumbent cells, some
›T
others with upright and squared cells.

Pith
Pith shape roundish. Cells dimorphic.
Cell content present (dark staining
substances). With prismatic crystals
Pistacia atlantica Desf.

and crystal druses. Pits in transverse


and in longitudinal cell walls. Vascu-
lar bundles clearly separate from one
other. Tracheary elements of metax-
ylem in distinct radial rows.

›T ›T

104
Stem xylem: 1 3 7 9 9.1 11 13 22 25 31 36 41 49 52.3 56 58 60 61 69 70.2 78 96 97 105 106 107 115 130 136 136.1
Twig bark: B1 B4 B6 B7 B9 B12 B14 B15 B17 B19 B20 B25 B26 B31 B33
Pith: P1 P4 P5 P6 P6.2 P9 P9.1 P10.1 P12

Anacardiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by the
difference in vessel size between late-
wood and earlywood. Wood ring-po-
rous. Earlywood vessels predominantly
solitary, latewood vessels in short
radial multiples and in clusters. Ves-
sels in diagonal and or radial pattern.
Mean tangential diameter of earlywood
vessel lumina 50-100 μm, 40-100 ves-
sels per mm2. Tyloses and deposits
in heartwood vessels. Fibers thin- to
thick-walled. Tension wood present.
Axial parenchyma scanty paratracheal.
Rays 4-12 per mm.
›T ›T

Radial section
Simple perforation plates. Interves-
sel pits alternate, small (4-7 μm in
diameter). Vessel-ray pits rounded or
angular with large apertures. Helical
thickenings in vessel elements present.
Earlywood vessel element length 100-
200 μm. Vascular and/or vasicentric
tracheids present. Fibers with simple
to minutely bordered pits (libriform
fibers). Body ray cells procumbent
with 1 to 3 rows of upright and square
marginal cells. Prismatic crystals in ray
parenchyma cells, and in tyloses.

›T ›T

Tangential section
Gradual transition from uniseriate to 4
cells wide rays. Multiseriate rays with
radial canals with epithelial cells.
Pistacia atlantica Desf.

Wood basic density: 0,74 g/cm³

›T ›T

105
Pistacia lentiscus L.
Mastic Tree, Lentisk
Anacardiaceae

PLANT DESCRIPTION
Evergreen, aromatic shrub or small
tree up to 4 m high. Leaves compound,
leaflets leathery, dark green. It occurs
in all the Mediterranean countries, in-
digenous to Cyprus. Very common on
rocky sites, sand dunes, maquis and
pine forests (0-800 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Fibers and scle-
reids present, arranged in tangential
groups. Prismatic crystals present.
Secretory elements in ducts present.
Phellem consist of large and small
cells, distinct in polarized light.

Xylem
Vessels solitary or in short radial
multiples. Mean tangential diameter
of earlywood vessels 20-50 μm. Early-
wood vessel element length 200-500
μm. Rays exclusively uniseriate, with
procumbent, square and upright cells ›T

mixed throughout the ray.


Pith
Pith shape roundish. Heterogeneous
pith. Cells dimorphic. Cell content pre-
sent (dark staining substances). Thick-
walled parenchyma cells scattered in
the pith. Prismatic crystals present.
Pits of two distinct sizes in transverse
Pistacia lentiscus L.

and in longitudinal cell walls. Vascular


bundles clearly separate. Tracheary el-
ements of metaxylem in distinct radial
rows. Axial cells in regular rows (radial
section).

›T ›T

106
Stem xylem: 1 3 7 9 10 11 13 22 25 31 36 41 49 52.3 60 69 70.2 78 96 97 106 115 130 136 136.1
Twig bark: B1 B4 B9 B12 B15 B17 B20 B25 B26 B32 B33
Pith: P1 P3 P3.3 P4 P5 P6 P9 P9.1 P9.3 P10.1 P12 P13

Anacardiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
the difference in vessel size between
latewood and earlywood. Wood ring-
porous. Vessels in radial multiples of 4
or more elements, and in clusters. Ves-
sels in diagonal and or radial pattern.
Mean tangential diameter of earlywood
vessel lumina 50-100 μm, 40-100
vessels per mm2. Fibers thin- to thick-
walled. Tension wood present. Axial
parenchyma scanty paratracheal. Rays
4-12 per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in diame-
ter). Vessel-ray pits rounded or angular
with large apertures. Helical thicken-
ings in vessel elements present. Early-
wood vessel element length 100-200
μm. Vascular and/or vasicentric trac-
heids present. Body ray cells procum-
bent with one row of upright or square
marginal cells. Prismatic crystals in ray
parenchyma cells.

›T ›T

Tangential section
Rays uniseriate to 3 cells wide. Radial
canals with epithelial cells.
Pistacia lentiscus L.

Wood basic density: 0,78 g/cm³

›T ›T

107
Pistacia terebinthus L.
Terebinth
Anacardiaceae

PLANT DESCRIPTION
Deciduous, aromatic shrub or small
tree up to 6 m high. Leaves alternate
and compound. Fruit an obovate
drupe, at first red, bluish-green at
maturity. A Mediterranean species, in-
digenous to Cyprus. It occurs on rocky
mountainsides and in pine forests,
maquis and garigue (0-1500 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube present. Fibers
in groups, sclereids predominantly in
tangential rows. Prismatic crystals and
crystal druses present. Phellem homo-
geneous, distinct in polarized light.

Xylem
Vessels in two distinct diameter class-
es.

›T

Pith
Pith shape polygonal to roundish. Cell
dimorphic. With prismatic crystals
and crystal druses. Pits in transverse
and in longitudinal cell walls. Vascular
Pistacia terebinthus L.

bundles clearly separate. Axial cells in


regular rows (radial section).

›T ›T

108
Stem xylem: 1 3 9 11 13 21 25 32 36 40.2 50.1 52.3 56 60 62 70 70.2 78 86 97 109 116.2 130 136 136.1 141.1
Twig bark: B1 B9 B14 B15 B17 B20 B22 B31 B33
Pith: P1.1 P4 P6 P6.2 P9 P9.1 P10.1 P13

Anacardiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
the difference in vessel size between
latewood and earlywood. Wood ring-
porous. Earlywood vessels mostly
solitary, vessel clusters common in
latewood. Mean tangential diameter of
earlywood vessels 20-50 μm, 100-200
vessels per mm2. Tyloses common.
Fibers very thick-walled. Tension wood
present. Axial parenchyma scanty para-
tracheal. More than 20 rays per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits opposite, small (4-7 μm in di-
ameter). Vessel-ray pits with large
horizontal or vertical apertures. Helical
thickenings in vessel elements present.
Earlywood vessels length 100-200 μm.
Vascular and/or vasicentric tracheids
present. Fibers with distinctly bor-
dered pits (fiber tracheids). Rays with
procumbent, square and upright cells
mixed throughout the ray. Prismatic
crystals in axial and ray parenchyma
cells.

›T ›T

Tangential section
Ray width 1 to 3 cells. Radial canals
with epithelial cells.
Pistacia terebinthus L.

Wood basic density: 0,81 g/cm³

›T ›T

109
Rhus coriaria L.
Sumac
Anacardiaceae

PLANT DESCRIPTION
Deciduous shrub up to 3 m high. Twigs
exuding milky latex when broken. Fruit
a suborbicular drupe, reddish-brown
at maturity. It occurs throughout the
Mediterranean and eastward to Iran;
native to Cyprus. Common on rocky
mountainsides, pine forests, maquis
and garigue (500-1600 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Only some rays
become dilated. Sclereids present,
scattered or irregularly dispersed.
Crystal druses present. With secretory ›T

elements in ducts.

Xylem
Vessels clusters common. Rays exclu-
sively uniseriate. Rays with procum-
bent, square and upright cells mixed
throughout the ray.

›T

Pith
Pith shape round to star-shaped. Cells
dimorphic. Crystal druses present. Pits
in transverse and in longitudinal cell
walls (radial section). Vascular bundles
clearly separate. Tracheary elements
of metaxylem in distinct radial rows.
Axial cells in regular rows (radial sec-
tion).
Rhus coriaria L.

›T ›T

110
Stem xylem: 1 3 7 9 11 13 22 26 27 31 36 42 48 53.1 56 60 61 69 70 70.3 77 79 97 106 115 136.1
Twig bark: B1 B4 B6 B15 B18 B22 B26
Pith: P1 P4 P4.1 P4.2 P6.2 P9 P9.1 P10.1 P12 P13

Anacardiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
the difference in vessel size between
latewood and earlywood, and by a lack
of vessels in terminal latewood. Wood
ring-porous. Vessels solitary and in
small clusters. Vessels in diagonal and
radial pattern in latewood. Mean tan-
gential diameter of earlywood vessel
lumina 100-200 μm, 20-40 vessels per
mm2. Tyloses common. Fibers thick- to
very thick-walled. Radial flat marginal
fibers present. Axial parenchyma dif-
fuse in aggregates and vasicentric.
Rays 4-12 per mm.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, medium to large (7
μm to more than 10 μm in diameter).
Vessel-ray pits rounded or angular with
large apertures. Helical thickenings in
vessel elements present. Earlywood
vessel element length 200-500 μm.
Vascular and/or vasicentric tracheids
present. Fibers with simple to minutely
bordered pits (libriform fibers). Body
ray cells procumbent with one row of
upright and square marginal cells. Pris-
matic crystals in ray parenchyma cells.

›T ›T

Tangential section
Ray width predominantly 1 to 3 cells.
Rhus coriaria L.

Wood basic density: 0,51 g/cm³

›T ›T

111
Schinus molle L.
Pepper Tree
Anacardiaceae

PLANT DESCRIPTION
Evergreen tree up to 10 m high, with
pendulous branches. Leaves alternate,
compound. Fruit a globose drupe,
rosy-red at maturity. A native of South
America, cultivated in various Mediter-
ranean countries; exotic to Cyprus.
Common in roadside plantations and
gardens (0-600 m alt.).

Photo: Börner
TWIG ANATOMY
Bark
Collapsed sieve tubes present. Some
rays become slightly dilated. Scleren-
chyma cells in phloem and in cortex.
Fibers scattered or irregularly dis-
persed, sometimes grouped. Prismatic
crystals present. With secretory ele-
ments in ducts. Phellem homogeneous.

Xylem
Growth rings boundaries distinct only
along some rays. Rays 12-20 per mm.

›T

Pith
Pith shape roundish. Cells dimorphic.
Cell content present (dark staining
substances). Prismatic crystals present.
Pits in transverse and in longitudinal
cell walls. Vascular bundles clearly
separate from one another. Tracheary
elements of metaxylem in distinct
radial rows. Axial cells in regular rows
Schinus molle L.

(radial section).

›T ›T

112
Stem xylem: 1 5 10 11 13 22 24 32 36 40.2 50.1 53.1 58 60 61 65 69 70.2 78 96 97 105 109 115 136 136.1
Twig bark: B4 B6 B7 B9 B13 B14 B20 B25 B14.1
Pith: P1 P4 P5 P6 P9 P9.1 P10.1 P10.2 P12 P13

Anacardiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
zones with fibers of variable cell wall
thickness. Wood diffuse-porous.
Vessels commonly in radial multiples
of 4 or more and in clusters. Mean tan-
gential diameter of earlywood vessels
20-50 μm, 100-200 vessels per mm2.
Gums and other deposits in heartwood
vessels. Fibers thin- to thick-walled.
Tension wood present. Axial paren-
chyma scanty paratracheal. Rays 4-12
per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, minute (less than 4 μm
in diameter). Vessel-ray pits with large
horizontal or vertical apertures. Helical
thickenings in vessel elements present.
Earlywood vessel element length 200-
500 μm. Vascular and/or vasicentric
tracheids present. Fibers with simple
to minutely bordered pits (libriform fib-
ers). Septate fibers present. Uniseriate
ray cells upright and square, multiseri-
ate rays with procumbent, square and
upright cells mixed throughout the ray.
Prismatic crystals and dark staining
substances in ray parenchyma cells.
›T ›T

Tangential section
Rays uniseriate and up to 3 cells wide.
Schinus molle L.

Wood basic density: 0,50 g/cm³

›T ›T

113
Schinus terebinthifolia Raddi
Brazilian Pepper Tree, Christmas-Berry Tree
Anacardiaceae

PLANT DESCRIPTION
Evergreen tree up to 15 m high. Fruit
a globose drupe, rosy-red at maturity.
A native of South America; cultivated
in various Mediterranean countries,
exotic to Cyprus. Common in roadside
plantations and gardens (0-400 m alt.).

TWIG ANATOMY
Bark
Some rays become dilated. Scleren-
chyma cells in phloem and in cortex.
Fibers grouped. Sclereids scattered or
irregularly dispersed. Prismatic crystals
present. Secretory elements in ducts
in cortex. Cell content in parenchyma
cells in cortex. Phellem homogeneous,
distinct in polarized light.

Xylem
Vessels in radial multiples of 4 or
more. Rays predominantly uniseriate.
More than 20 rays per mm.
›T

Pith
Schinus terebinthifolia Raddi

Pith shape round. Cells dimorphic.


With prismatic crystals and crystal
druses. Intercellular canals with border
cells. Pits in transverse and in longi-
tudinal cell walls. Vascular bundles
clearly separate. Tracheary elements of
metaxylem in distinct radial rows.

›T ›T

114
Stem xylem: 1 5 9 9.1 13 22 24 32 36 40.2 49 53.1 58 61 65 69 70.2 78 96 97 106 107 115 130 136 136.1
Twig bark: B6 B7 B9 B14 B15 B18 B20 B25 B26 B28 B31 B33
Pith: P1 P4 P6 P6.2 P8 P9 P9.1 P10.1 P12

Anacardiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood diffuse-porous. Vessels
solitary and in radial multiples of 2 to
6 elements. Mean tangential diameter
of earlywood vessels 20-50 μm, 40-
100 vessels per mm2. Gums and other
deposits in heartwood vessels. Fibers
thin- to thick-walled. Tension wood
present. Axial parenchyma scanty para-
tracheal. Rays 4-12 per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, minute (less than 4 μm
in diameter). Vessel-ray pits with large
horizontal or vertical apertures. Helical
thickenings in vessel elements present.
Earlywood vessel element length 200-
500 μm. Fibers with simple to minutely
bordered pits (libriform fibers). Septate
fibers present. Body ray cells procum-
bent with one row of upright marginal
cells. Prismatic crystals in ray paren-
chyma cells.

›T ›T

Tangential section Schinus terebinthifolia Raddi


Rays uniseriate and up to 3 cells wide.
Intercellular radial canals with border
cells.

Wood basic density: 0,56 g/cm³

›T ›T

115
Nerium oleander L.
Oleander
Apocynaceae

PLANT DESCRIPTION
Evergreen, lactiferous shrub 2-4 m
high. Leaves in whorls of 3 or oppo-
site, thick and coriaceous. Flowers
dark to light pink or white. Widespread
in the Mediterranean region. Native of
Cyprus, very common along streams of
the island and widely planted for orna-
ment (0-900 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube present. Sclereids
scattered or irregularly dispersed. Fib-
ers with an unlignified innermost layer
in groups. Prismatic crystals present.
With laticifers, secretory elements,
oil ducts or mucilage ducts in cortex.
Phellem homogeneous.

Xylem
Vessels mostly in radial multiples of
4 or more. Earlywood vessel element
length more than 500 μm. Rays exclu-
sively uniseriate. More than 20 rays
per mm.
›T

Pith
Pith not visible in polarized light. Pith
shape triangular. Medullary sheath pre-
sent. Heterogenous pith. Sclereids pre-
sent. Cell content present (dark stain-
ing substances). With prismatic crystals
and crystal druses. Pits in transverse
Nerium oleander L.

and in longitudinal cell walls. Vascular


bundles at the periphery of the pith.

›T ›T

116
Stem xylem: 1 5 9 9.1 10 13 22 24 30 40.2 49 53.1 61 65 69 76 86 96.1 97 100 109 116.1 136 136.1 200
Twig bark: B1 B4.1 B10 B15 B18 B20 B25 B31
Pith: P1.1 P1.2 P2 P3 P3.1 P5 P6 P6.2 P9 P9.1 P10

Apocynaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
zones with fibers of variable cell wall
thickness. Wood diffuse-porous. Ves-
sels solitary or in radial multiples of 2
to 4 or more. Mean tangential diameter
of earlywood vessels 20-50 μm, 40-100
vessels per mm2. Fibers thin- to thick-
walled. Apotracheal parenchyma dif-
fuse and in narrow bands or lines up to
three cells wide. Rays 12-20 per mm.

›T ›T

Radial section
Simple perforation plates. Interves-
sel pits alternate, minute (less than 4
μm in diameter). Vessel-ray pits with
distinct borders, similar to interves-
sel pits in size and shape throughout
the ray cell. Earlywood vessel element
length 200-500 μm. Fibers with simple
to minutely bordered pits (libriform fib-
ers). Septate fibers present. Rays with
procumbent, square and upright cells
mixed throughout the ray. Few pris-
matic crystals in ray parenchyma cells.

›T ›T

Tangential section
Rays uniseriate to 3 cells wide. Rays
with multiseriate portions as wide as
uniseriate portions.
Nerium oleander L.

Wood basic density: 0,41 g/cm³

›T ›T

117
Vinca major L.
Grater Periwinkle
Apocynaceae

PLANT DESCRIPTION
Evergreen creeping subshrub, with
sterile shoots, up to 1 m long, and
shorter suberect flowering stems.
Leaves opposite, broadly ovate. Native
of western and central Europe and the
Mediterranean countries, naturalized in
Cyprus. It is found in gardens and fal-
low land (0-1600 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclereids in tan-
gential rows, scattered or irregularly
dispersed. Fibers with an unlignified
innermost layer in tangentially ar-
ranged groups. Cortex homogeneous.
Epidermis distinct in polarized light.

Xylem
As stem wood.

›T

Pith
Pith not visible in polarized light. Pith
shape round. Medullary sheath pre-
sent. Unlignified cells present. Pits
in transverse and in longitudinal cell
walls. Vascular bundles at the periph-
ery of the pith.
Vinca major L.

›T ›T

118
Stem xylem: 2.1 5 9 9.1 13 22 25 36 40.2 50.2 53.1 61 65 69 75 96 100.1 105 116.1
Twig bark: B1 B4 B15 B17 B18 B31 B33
Pith: P1 P2 P3.4 P9 P9.1 P10

Apocynaceae
STEM ANATOMY
Transverse section
Only one ring in the observed sample.
Wood diffuse-porous. Vessels solitary
and in radial multiples of 2 to 4 ele-
ments. Mean tangential diameter of
vessels 20-50 μm, more than 200
vessels per mm2. Fibers thin- to thick-
walled. Axial parenchyma absent,
extremely rare or not to recognizable.
Rays 12-20 per mm.

›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Helical thickenings in ves-
sel elements present. Vessel element
length 200-500 μm. Fibers with simple
to minutely bordered pits (libriform
fibers). Septate fibers present. All ray
cells upright.

›T ›T

Tangential section
Rays exclusively uniseriate. Rays con-
fluent with ground tissue.
Vinca major L.

Wood basic density: 0,27 g/cm³

›T

119
Ilex aquifolium L.
Holly
Aquifoliaceae

PLANT DESCRIPTION
Evergreen shrub up to 10 m high, with
dense spherical or ovoid crown. Leaves
with spinose or cuspidate margins,
leathery. Flowers small and white. Fruit
a globose, red drupe. Exotic to Cyprus,
native of many European and Mediter-
ranean countries and eastward to Iran.

TWIG ANATOMY
Bark
Groups of sieve tube present. Fibers
scattered or irregularly dispersed and
grouped. Sclereids in groups. Prismatic
crystals and crystal druses present.
Phloem uniform. Cortex homogene-
ous. Epidermis distinct in polarized
light. Lignified cells in epidermis.

Xylem
As stem wood.

›T

Pith
Pith shape round. Thick-walled paren-
chyma cells present. Cells dimorphic.
Cell content present (dark staining
substances). Prismatic crystals and
crystal sand present. Pits in transverse
and in longitudinal cell walls. Vascular ›T
bundles clearly separate. Tracheary el-
Ilex aquifolium L.

ements of metaxylem in distinct radial


rows.

›T

120
Stem xylem: 1 4 5 7 10 14 16 17 21 26 30 36 40.2 50.2 53.1 62 69 76 98 102 103 106 107 115
Twig bark: B1 B9 B13 B14 B15 B19 B20 B22 B8 B31 B33 B34
Pith: P1 P3.3 P4 P5 P6 P6.4 P9 P9.1 P10.1 P12

Aquifoliaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted latewood fibers. Wood
diffuse-porous to semi-ring-porous.
Vessels in radial pattern. Vessels in
radial multiples of 4 or more elements
common. Mean tangential diameter
of earlywood vessels 20-50 μm. More
than 200 vessels per mm2. Fibers thin-
to thick-walled. Axial parenchyma dif-
fuse. Rays 4-12 per mm.

›T ›T

Radial section
Scalariform perforation plates with
20-40 bars. Inter-vessel pits oppo-
site, medium (7-10 μm in diameter).
Vessel-ray pits with distinct borders,
similar to intervessel pits in size and
shape throughout the ray cell. Helical
thickenings in vessel elements present.
Earlywood vessel element length 200-
500 μm. Fibers with distinctly bordered
pits (fiber tracheids). Body ray cells
procumbent with 1-3 rows of upright
and square marginal cells.

›T ›T

Tangential section
Larger rays uniseriate and up to 7 seri-
ate. Ray height more than 1 mm.
Ilex aquifolium L.

Wood basic density: 0,78 g/cm³

›T ›T

121
Hedera helix L. = Hedera helix L. subsp. poetarum Nyman
Ivy
Araliaceae

PLANT DESCRIPTION
Evergreen climber with aerial roots.
Leaves alternate, simple; lamina of
leaves on sterile shoots cordate or del-
toid, leaves on flowering shoots lamina
ovate to broadly lanceolate. Native to
Cyprus, growing in moist sites (100-
1600 m alt.). It also occurs in temper-
ate Europe and Asia.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclereids in tan-
gential rows, scattered or irregularly
dispersed. Phellem homogeneous,
distinct in polarized light.

Xylem
Growth rings clearly demarcated only
along some radii. Mean tangential di-
ameter of earlywood vessels 20-50 μm,
earlywood vessels length more than
500 μm. Rays 12-20 per mm.

›T

Pith
Pith not visible in polarized light.
Pith shape round. Medullary sheath
present. Cells dimorphic. Unlignified
cells present. Pits in transverse and in
longitudinal cell walls. Vascular bun-
dles clearly separate.
Hedera helix L.

›T ›T

122
Stem xylem: 1 3 4 11 13 20 22 26 31 41 50.1 53.1 60 61 65 67 69 76 78 96 99 102 103 104 109 115
Twig bark: B1 B4 B15 B17 B18 B31 B33
Pith: P1 P2 P3.4 P4 P4.1 P9 P9.1 P10.1

Araliaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by a
lack of vessels in terminal latewood.
Wood ring-porous to semi-ring-porous.
Vessel clusters common. Mean tan-
gential diameter of earlywood vessel
lumina 50-100 μm, 100-200 vessels
per mm2. Parenchyma-like fiber bands
alternating with ordinary fibers. Fib-
ers thin- to thick-walled. Apotracheal
parenchyma diffuse to scanty paratra-
cheal. Rays 4-12 per mm.

›T ›T

Radial section
Simple perforation plates. Few inter-
vessel pits scalariform. Inter-vessel pits
alternate, medium (7-10 μm in diam-
eter). Vessel-ray pits rounded or an-
gular with large apertures. Earlywood
vessel element length 200-500 μm.
Vascular and/or vasicentric tracheids
present. Fibers with simple to minutely
bordered pits (libriform fibers). Sep-
tate fibers present. Rays with procum-
bent, square and upright cells mixed
throughout the ray.

›T ›T

Tangential section
Rays of two distinct sizes: uniseriate
and more than 10 seriate. Larger rays
height more than 1 mm.
Hedera helix L.

Wood basic density: 0,36 g/cm³

›T ›T

123
Aristolochia sempervirens L.
Climbing Birthwort, Dutchman’s Pipe
Aristolochiaceae

PLANT DESCRIPTION
Evergreen climber up to 3 m high, with
a thick root stock. Leaves cordate with
acute apex, coriaceous. Flowers tubu-
lar, strongly curved. Indigenous to Cy-
prus growing among trees and shrubs
usually in moist situations (0-1200 m
alt.). Native also to Algeria, Sicily east-
ward to Greece and Lebanon.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Fibers arranged in
wide tangential bands in cortex. Scle-
reids scattered or irregularly dispersed.
Cortex consist of a belt of thick-walled
fibers and unlignified parenchyma
cells. Epidermis distinct in polarized
light. Lignified cells in epidermis.

Xylem
As stem wood.

›T

Pith
Aristolochia sempervirens L.

Unlignified cells present. Vascular


bundles not distinct. Pith collapsed in
older twigs and in stem.

›T

124
Stem xylem: 1 3 9 13 22 26 39.1 40.2 45 50.2 52.3 62 69 78 99 100.2 109 114
Twig bark: B1 B4 B9 B12 B15 B18 B14.1
Pith: P3.4 P10.2

Aristolochiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
the difference in vessel size between
latewood and earlywood. Wood ring-
porous. Vessels predominantly solitary.
Vessels cell wall thick (more than 2
μm). Mean tangential diameter of early-
wood vessels 20-50 μm. Vessels of two
distinct diameter classes. More than
200 vessels per mm2. Fibers thin- to
thick-walled. Axial parenchyma scanty
paratracheal. Less than 4 rays per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, medium (7-10 μm in
diameter). Earlywood vessel element
length 100-200 μm. Fibers with dis-
tinctly bordered pits (fiber tracheids).
Rays with procumbent, square and
upright cells mixed throughout the ray.

›T ›T

Tangential section Aristolochia sempervirens L.


Larger rays commonly more than 10
seriate. Rays not visible in polarized
light.

Wood basic density: 0,29 g/cm³

›T ›T

125
Cyprinia gracilis (Boiss.) Browicz
Cyprinia
Asclepiadaceae

PLANT DESCRIPTION
Deciduous, slender climber reaching
more than 8 m in length. Shoots with
milky latex. Leaves opposite, simple,
leathery. The genus Cyprinia includes
a single species, which is indigenous
to Cyprus and southern Turkey. On
Cyprus it is rather uncommon but not
rare (0-1100 m alt.).

TWIG ANATOMY
Bark
Collapsed sieve tubes present. Scle-
reids scattered or irregularly dispersed
and in groups. Prismatic crystals pre-
sent. With laticifers, secretory elements
in ducts.

Xylem
First growth ring consist of 4-6 con-
secutive radially arranged small vessels
and fibers, with few 50-100 μm diam-
eter vessels.

›T
Cyprinia gracilis (Boiss.) Browicz

Pith
Pith not visible in polarized light. Pith
shape roundish. Medullary sheath pre-
sent. Cells dimorphic. Unlignified cells
present. Large vascular bundles in the
pith. Tracheary elements of metaxylem
in distinct radial rows.

›T ›T

126
Stem xylem: 1 3 9 9.1 13 22 25 30 42 48 53.1 62 69 70.3 78 84 96 109 116.2
Twig bark: B4 B15 B18 B19 B20 B25 B27
Pith: P1.1 P1 P3.4 P4 P10 P12

Asclepiadaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
the difference in vessel size between
latewood and earlywood and by radi-
ally flatted latewood fibers. Wood ring-
porous. Earlywood vessels predomi-
nantly solitary, latewood vessels in
short radial multiples common. Mean
tangential diameter of earlywood ves-
sel lumina 100-200 μm, 20-40 vessels
per mm2. Fibers thin- to thick-walled.
Radial flat marginal fibers. Axial paren-
chyma scanty paratracheal and unilat-
eral paratracheal. More than 20 rays
per mm.
›T ›T

Radial section
Simple perforation plates. Interves-
sel pits alternate, small (4-7 μm in
diameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 200-
500 μm. Fibers with distinctly bor-
dered pits (fiber tracheids). Rays with
procumbent, square and upright cells
throughout the ray.

›T ›T

Tangential section Cyprinia gracilis (Boiss.) Browicz


Rays exclusively uniseriate.

Wood basic density: 0,36 g/cm³

›T ›T

127
Achillea cretica L.
Asteraceae

PLANT DESCRIPTION
Much branched, neatly rounded sub-
shrub up to 50 cm high, with densely
white-tomentose shoots. Leaves nu-
merous, linear (10-25x2-3 mm). Indig-
enous to Crete, the Aegean islands and
southwest Turkey. A rare indigenous
species on Cyprus, occurring on rocky
slopes near the sea (0-150 m).

TWIG ANATOMY
Bark
Groups of sieve tube present. Larger
rays become dilated. Large groups of
fibers containing scattered sieve tubes
and parenchyma cells between rays.
Crystal sand present. Phellem consist
of thin-walled and irregularly shaped
cells.

Xylem
First growth ring with few small ves-
sels and very thick-walled fibers. Rays
predominantly uniseriate.

›T

Pith
Pith shape round. Cells dimorphic.
Crystal sand present. Vascular bundles
clearly separate.
Achillea cretica L.

›T ›T

128
Stem xylem: 1 5 9.1 11 13 22 25 30 40.2 45 50.2 52.3 58 60 61 69 78 89 96 97 98 102 105 114 136 136.1
Twig bark: B1 B6 B9 B12 B14 B23
Pith: P1 P4 P6.4 P10.1

Asteraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted latewood fibers, and
by initial marginal parenchyma cells.
Wood diffuse-porous. Vessels in short
radial multiples and in clusters. Mean
tangential diameter of earlywood ves-
sels 20-50 μm. More than 200 vessels
per mm2. Vessels of two distinct diam-
eter classes. Gums and other deposits
in heartwood vessels. Fibers thin- to
thick-walled. Axial parenchyma scanty
paratracheal. Apotracheal parenchyma
in marginal or in seemingly marginal
bands. Less than 4 rays per mm.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 100-
200 μm. Vascular and/or vasicentric
tracheids present. Fibers with simple
to minutely bordered pits (libriform
fibers). All ray cells upright and square.
Elongated crystals in ray cells.

›T ›T

Tangential section
Rays from uniseriate to 10 seriate.
Larger rays more than 1 mm high.
Achillea cretica L.

Wood basic density: 0,42 g/cm³

›T ›T

129
Achillea maritima (L.) Ehrend. et Y.P. Guo subsp. maritima
= Otanthus maritimus (L.) Hoffmanns. et Link subsp. maritimus
Asteraceae

PLANT DESCRIPTION
Subshrub up to 40 cm high. Stems
erect or sprawling, often much-
branched. Leaves oblong, sessile,
densely white-tomentose. Indigenous
to south-western Europe and the
Mediterranean region. A rare native of
Cyprus occurring on sand dunes and
sandy seashores (0 m alt.).

TWIG ANATOMY
Achillea maritima (L.) Ehrend. et Y.P. Guo subsp. maritima

Bark
Groups of sieve tube in radial rows.
Fibers in tangential groups and scat-
tered or irregularly dispersed. Crystal
sand present. Cortex consist of large
and small thin-walled parenchyma
cells.

Xylem
As stem wood.

›T

Pith
Pith shape round. Unlignified cells
present. Heterogeneous pith. Pits in
transverse and in longitudinal cell
walls. Vascular bundles clearly sepa-
rate. Axial cells in regular rows (radial
section).

›T ›T

130
Stem xylem: 1.1 5 9.1 13 22 25 30 40.1 40.2 45 50.1 52.3 60 61 65 67 69 78 96 98 102 103 105 115
Twig bark: B1 B3 B9 B12 B7.3
Pith: P1 P3.4 P9 P10.1 P13

Asteraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct only
along some radii. Wood diffuse-porous.
Vessels solitary or in short radial mul-
tiples. Mean tangential diameter of
earlywood vessels 20-50 μm, 100-200
vessels per mm2. Vessels of two dis-
tinct diameter classes. Fibers thin- to
thick-walled. Axial parenchyma scanty
paratracheal. Rays 4-12 per mm.

›T ›T

Radial section

Achillea maritima (L.) Ehrend. et Y.P. Guo subsp. maritima


Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 100-
200 μm. Vascular and/or vasicentric
tracheids present. Fibers with simple
to minutely bordered pits (libriform
fibers). Septate fibers present. Paren-
chyma-like fiber bands alternating with
ordinary fibers. All ray cells upright
and square.

›T ›T

Tangential section
Rays of two distinct sizes: uniseri-
ate and up to 10 seriate. Larger rays
height more than 1 mm.

Wood basic density: 0,33 g/cm³

›T ›T

131
Ambrosia maritima L.
Damaseisa, Sea Ambrosia
Asteraceae

PLANT DESCRIPTION
Aromatic perennial herb up to 1 m
high. Male capitula in leafless, terminal
racemes; female inflorescence in the
axils of the uppermost leaves. It occurs
in the Mediterranean region; indig-
enous to Cyprus, occurring on sandy
seashores or muddy canal banks (0-20
m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube present. Some
rays become slightly dilated. Few fib-
ers scattered or irregularly dispersed.
Large groups of fibers present. Pris-
matic crystals and crystal sand pre-
sent. Cortex consist of unlignified
parenchyma cells and groups of thick-
walled fibers. Epidermis distinct in
polarized light.

Xylem
Vessels predominantly solitary or in
short radial multiples. Axial parenchy-
ma scanty paratracheal.
›T

Pith
Pith not visible in polarized light. Pith
shape round. Unlignified cells present.
Prismatic crystals and crystal druses
present. Vascular bundles clearly sepa-
Ambrosia maritima L.

rate. Tracheary elements of metaxylem


in distinct radial rows. Axial cells in
regular rows (radial section).

›T ›T

132
Stem xylem: 1 5 10 11 13 22 25 30 40.2 45 50.2 53.1 56 60 61 69 78 89 89.1 96 98 102 103 105 114
Twig bark: B1 B6 B13 B14 B20 B23 B33
Pith: P1 P3.4 P6 P6.2 P10.1 P12 P13

Asteraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
initial marginal parenchyma bands.
Wood diffuse-porous. Vessels in radial
multiples of 4 or more elements, and
in clusters. Mean tangential diameter
of earlywood vessels 20-50 μm. More
than 200 vessels per mm2. Vessels of
two distinct diameter classes. Tyloses
with thin walls common. Fibers thin- to
thick-walled. Parenchyma scanty para-
tracheal. Apotracheal parenchyma cells
in marginal or in seemingly marginal
rows, thin-walled, dark in polarized
light. Less than 4 rays per mm.
›T ›T

Radial section
Simple perforation plates. Interves-
sel pits alternate, small (4-7 μm in
diameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 200-
500 μm. Vascular and/or vasicentric
tracheids present. Fibers with simple
to minutely bordered pits (libriform fib-
ers). All ray cells upright.

›T ›T

Tangential section
Rays of two distinct sizes: uniseri-
ate and up to 10 seriate. Larger rays
height more than 1 mm.
Ambrosia maritima L.

Wood basic density: 0,31 g/cm³

›T ›T

133
Artemisia arborescens L.
Shrubby Wormwood
Asteraceae

PLANT DESCRIPTION
Aromatic shrub up to 1 m high. Stems
and leaves covered with silvery hairs.
Widespread in the Mediterranean re-
gion. Naturalized on Cyprus, found as
a relic or escape of cultivation in hedg-
es, dry stone walls and field borders
(50-1400 m alt.). It is cultivated also as
a medical plant.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Some rays become
dilated. Sclerenchyma cells in phloem
and in cortex. Fibers in groups. Scle-
reids scattered or irregularly dispersed
and in groups. Crystal sand present.
Ducts with border cells in cortex. Phel-
lem homogeneous, distinct in polar-
ized light.

Xylem
Mean tangential diameter of earlywood
vessels less than 30 μm. All ray cells
upright and square. In the first ring
›T
very thick-walled fibers present.

Pith
Pith shape round. With crystal druses
and crystal sand present. Pits of two
Artemisia arborescens L.

distinct size, grouped both in trans-


verse and in longitudinal cell walls.
Vascular bundles clearly separate.
Tracheary elements of metaxylem in
distinct radial rows.

›T ›T

134
Stem xylem: 1 4 5 9 9.1 11 13 22 25 30 36 40.2 45 50.2 52.3 60 61 69 78 96 97 105 109 115
Twig bark: B1 B4 B6 B7 B9 B14 B15 B18 B19 B23 B25 B26 B31 B14.3
Pith: P1 P6.2 P6.4 P9 P9.1 P9.2 P9.3 P10.1 P12

Asteraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
initial marginal parenchyma cells, and
by zones with fibers of variable cell
wall thickness. Wood diffuse-porous to
semi-ring-porous. Vessels solitary or in
short radial multiples, and in clusters.
Mean tangential diameter of earlywood
vessels 20-50 μm. More than 200 ves-
sels per mm2. Vessels of two distinct
diameter classes. Fibers thin- to thick-
walled. Axial parenchyma scanty para-
tracheal. Rays 4-12 per mm.

›T ›T

Radial section
Simple perforation plates. Intervessel
pits alternate, small (4-7 μm in diame-
ter). Vessel-ray pits with distinct bor-
ders, similar to intervessel pits in size
and shape throughout the ray cell. Fine
helical thickenings in vessel elements
present. Earlywood vessel element
length 100-200 μm. Vascular and/
or vasicentric tracheids present. Fib-
ers with simple to minutely bordered
pits (libriform fibers). Most uniseriate
rays composed by upright and square
cells. Multiseriate rays with square, up-
right and few procumbent cells mixed
throughout the ray.
›T ›T

Tangential section
Rays uniseriate to 3 cells wide.
Artemisia arborescens L.

Wood basic density: 0,54 g/cm³

›T ›T

135
Centaurea akamantis T.Georgiades et Hadjik.
Akamas Centaury
Asteraceae

PLANT DESCRIPTION
Subshrub with hanging or suberect,
densely hairy shoots, up to 60 cm
long. A very rare endemic to Cyprus,
confined in two neibouring gorges in
the Akamas peninsula (50-100 m alt.).
Centaurea akamantis in strictly pro-
tected.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Some rays become
slightly dilated. Fibers grouped. Crystal
sand present. Cortex consist of thin-
walled parenchyma cells and groups of
thick-walled fibers.
Centaurea akamantis T.Georgiades et Hadjik.

Xylem
Only one ring in the observed sample.
Axial parenchyma absent or extremely
rare or not to recognizable.

›T

Pith
Pith shape roundish to pentagonal.
Cells dimorphic. Pits in transverse
and in longitudinal cell walls. Vascular
bundles clearly separate. Axial cells in
regular rows (radial section).
›T

›T

136
Stem xylem: 1 1.1 5 11 13 22 25 30 39.1 40.2 45 50.1 52.3 60 61 65 69 70 78 79 96 98 102 103 105 114
Twig bark: B1 B4 B6 B14 B23 B14.1
Pith: P1.1 P4 P9 P9.1 P10.1 P13

Asteraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct only
along some radii by the difference
in vessel size between latewood and
earlywood and by the presence of mar-
ginal parenchyma cells. Wood diffuse-
porous. Vessels predominantly in
clusters. Vessels cell wall thick (more
than 2 μm). Mean tangential diameter
of earlywood vessels 20-50 μm, 100-
200 per mm2. Vessels in two distinct
diameter classes. Fibers thick- to very
thick-walled. Axial parenchyma scanty
paratracheal to vasicentric. Less than 4
rays per mm.
›T ›T

Radial section
Simple perforation plates. Interves-
sel pits alternate, small (4-7 μm in
diameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 100-
200 μm. Vascular and/or vasicentric
tracheids present. Fibers with simple

Centaurea akamantis T.Georgiades et Hadjik.


to minutely bordered pits (libriform
fibers). Septate fibers present. All ray
cells upright and square.

›T ›T

Tangential section
Rays of two distinct sizes: uniseri-
ate and up to 10 seriate. Larger rays
height more than 1 mm.

Wood basic density: 0,30 g/cm³

›T ›T

137
Cichorium spinosum L.
Spiny Chicory
Asteraceae

PLANT DESCRIPTION
Intricately branched subshrub up to
50 cm high, with repeatedly branched
stems. Widely distributed throughout
the Mediterranean basin from Spain
to Cyprus. Indigenous to the island,
occurring on rock-crevices by the sea,
and sometimes on sandy coastal fields
(0-200 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube present. Col-
lapsed sieve tubes in radial rows. Crys-
tal sand present. Phellem homogene-
ous. Cortex consist of large and small
unlignified parenchyma cells.

Xylem
As stem wood.

›T

Pith
Pith not visible in polarized light. Pith
shape polygonal to round. Heteroge-
neous pith. Unlignified cells present.
Pits in transverse and in longitudinal
Cichorium spinosum L.

cell walls. Vascular bundles clearly


separate to not distinct. Tracheary ele-
ments of metaxylem in distinct radial
rows. Axial cells in regular rows (radial
section).

›T ›T

138
Stem xylem: 1 5 10 11 13 22 25 30 40.2 45 50.2 52.3 60 61 78 89 89.1 96 97 102 103 105 116.2
Twig bark: B1 B4 B23 B31
Pith: P1.1 P1.1 P3 P3.4 P9 P9.1 P10.1 P10.2 P12 P13

Asteraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ini-
tial marginal parenchyma bands. Wood
diffuse-porous. Vessels in radial mul-
tiples of 4 or more elements, and in
clusters. Mean tangential diameter of
earlywood vessels 20-50 μm. Vessels
of two distinct diameter classes. More
than 200 vessels per mm2. Axial paren-
chyma scanty paratracheal. Apotrache-
al parenchyma cells in marginal or in
seemingly marginal bands, thin-walled,
dark in polarized light. More than 20
rays per mm.
›T ›T

Radial section
Simple perforation plates. Interves-
sel pits alternate, small (4-7 μm in
diameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 100-
200 μm. Vascular and/or vasicentric
tracheids present. Fibers with simple
to minutely bordered pits (libriform fib-
ers). All ray cells upright and square.

›T ›T

Tangential section
Rays of two distinct sizes: uniseriate
and up to 3 cells wide, the latter more
than 1 mm height.
Cichorium spinosum L.

Wood basic density: 0,33 g/cm³

›T ›T

139
Dittrichia viscosa (L.) Greuter subsp. angustifolia (Bég.) Greuter
= Inula viscosa (L.) Aiton
Asteraceae

PLANT DESCRIPTION
Viscid-glutinous, aromatic subshrub
up to 1,5 m high. Indigenous to the
Mediterranean region and the Atlan-
tic islands. Very common on Cyprus,
growing on disturbed grounds, along
roadsides, hillsides, often in moist
sites and near springs or rivers (0-
1600 m alt.).
Dittrichia viscosa (L.) Greuter subsp. angustifolia (Bég.) Greuter

TWIG ANATOMY
Bark
Groups of sieve tube present. Some
rays become dilated. Fibers in tangen-
tial rows, scattered or irregularly dis-
persed. Fibers in large groups. Acicular
crystals and crystal sand present. Cor-
tex consist of thin-walled parenchyma
cells and groups of thick-walled fibers.
Phellem homogeneous.

Xylem
Vessels predominantly in short radial
multiples. Mean tangential diameter of
earlywood vessel lumina 50-100 μm.
More than 200 vessels per mm2.
›T

Pith
Pith shape round. Cells dimorphic.
Acicular crystals present. Pits grouped
in transverse and in longitudinal cell
walls. Vascular bundles clearly sepa-
rate. Tracheary elements of metaxylem
in distinct radial rows. Axial cells in
regular rows (radial section).

›T ›T

140
Stem xylem: 1 4 5 9 13 22 25 30 40.2 45 50.2 53.1 60 61 69 78 96 98 102 103 105 114
Twig bark: B1 B6 B12 B13 B14 B21 B23 B14.1
Pith: P1 P4 P6.1 P9 P9.1 P9.2 P10.1 P12 P13

Asteraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
the difference in vessel size between
latewood and earlywood and by ra-
dially flatted latewood fibers. Wood
semi-ring to diffuse-porous. Vessels
predominantly solitary. Mean tangen-
tial diameter of earlywood vessels
20-50 μm. More than 200 vessels per
mm2. Vessels of two distinct diameter
classes. Fibers thin- to thick-walled.
Axial parenchyma scanty paratracheal.
Less than 4 rays per mm.

Dittrichia viscosa (L.) Greuter subsp. angustifolia (Bég.) Greuter


›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 200-
500 μm. Vascular and/or vasicentric
tracheids present. Fibers with simple
to minutely bordered pits (libriform fib-
ers). All ray cells upright and square.

›T ›T

Tangential section
Rays of two distinct sizes: uniseriate
and 4 to 10 seriate. Larger ray height
more than 1 mm.

Wood basic density: 0,48 g/cm³

›T ›T

141
Helichrysum italicum (Roth) G.Don
Everlasting
Asteraceae

PLANT DESCRIPTION
Aromatic shrub up to 80 cm high, with
a characteristic curry smell. Flowering
stems erect, ascending of flexuous.
Leaves linear, hairy. Widely distributed
in the Mediterranean region. Indig-
enous to Cyprus, occurring on rocky,
usually igneous, mountainsides (500-
1800 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Some rays become
slightly dilated. Sclerenchyma cells in
phloem and in cortex. Fibers arranged
in large and dense groups with scat-
tered sieve tubes.

Xylem
As stem wood.

›T
Helichrysum italicum (Roth) Don

Pith
Pith shape round. Heterogeneous pith.
Unlignified cells present. Pits in trans-
verse and in longitudinal cell walls.
Vascular bundles clearly separate.
Tracheary elements of metaxylem in
distinct radial rows. Axial cells in regu-
lar rows (radial section).

›T ›T

142
Stem xylem: 1 4 5 9.1 11 13 22 25 30 36 40.2 45 50.1 52.3 58 60 61 69 70.3 78 89 89.1 96 98 102 105 115 124
Twig bark: B1 B4 B6 B7 B12 B5.4
Pith: P1 P3 P3.4 P9 P9.1 P10.1 P12 P13

Asteraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers and by initial marginal parenchy-
ma cells. Wood semi-ring-porous to dif-
fuse-porous. Vessels in radial multiples
of 2 to 4 elements, and in clusters.
Mean tangential diameter of earlywood
vessels 20-50 μm, 100-200 vessels per
mm2. Vessels of two distinct diameter
classes. Deposits in heartwood vessels.
Fibers thin- to thick-walled. Parenchy-
ma scanty paratracheal. Apotracheal
parenchyma in marginal or in seem-
ingly marginal bands, dark in polarized
›T ›T
light. Rays 4-12 per mm.

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Helical thickenings in vessel elements
present. Earlywood vessel element
length 100-200 μm. Vascular and/or
vasicentric tracheids present. Fibers
with simple to minutely bordered pits
(libriform fibers). All ray cells upright
and square.

›T ›T

Tangential section Helichrysum italicum (Roth) Don


Rays uniseriate to 10 seriate. Larger
rays more that 1 mm high. Oil and/
or mucilage cells associated with ray
parenchyma.

Wood basic density: 0,64 g/cm³

›T ›T

143
Helichrysum stoechas subsp. barrelieri (Ten.) Nyman
= Helichrysum conglobatum (Viv.) Steud.
Asteraceae

PLANT DESCRIPTION
Subshrub up to 80 cm high with arach-
noid, erect shoots. Leaves woolly hairy.
A common indigenous species on Cy-
prus, growing in garigue, maquis and
rocky hillsides (0–700 m alt.). Widely
distributed in the Mediterranean re-
gion.

TWIG ANATOMY
Bark
Collapsed sieve tubes present. Some
rays become dilated. Fibers in large
Helichrysum stoechas subsp. barrelieri (Ten.) Nyman

tangentially elongated groups, with


few scattered sieve tubes. Anatomy dif-
ficult to interpret.

Xylem
As stem wood.

›T

Pith
Pith shape round. Cells dimorphic. Pits
in transverse and in longitudinal cell
walls. Vascular bundles clearly sepa-
rate. Axial cells in regular rows (radial
section).

›T ›T

144
Stem xylem: 1 5 7 9.1 11 13 21 22 24 30 36 39.1 40.2 45 50.1 52.3 61 67 69 70.3 78 89 96 98 105 114
Twig bark: B4 B6 B9 B5.4
Pith: P1 P4 P4.1 P9 P9.1 P10.1 P13

Asteraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood diffuse-porous. Vessels in
radial pattern. Vessels in short radial
multiples and in clusters. Vessels cell
wall thick (more than 2 μm). Mean
tangential diameter of earlywood ves-
sels 20-50 μm, 100-200 vessels per
mm2. Vessels of two distinct diameter
classes. Parenchyma-like fiber bands
alternating with ordinary fibers. Fib-
ers thin- to thick-walled. Radial flat
marginal fibers. Parenchyma scanty
›T ›T
paratracheal. Apotracheal parenchyma
in marginal or in seemingly marginal
bands. Less than 4 rays per mm.
Radial section
Simple perforation plates. Inter-vessel
pits opposite and alternate, minute
(less than 4 μm in diameter). Vessel-

Helichrysum stoechas subsp. barrelieri (Ten.) Nyman


ray pits with distinct borders, similar
to intervessel pits in size and shape
throughout the ray cell. Helical thick-
enings in vessel elements present.
Earlywood vessels length 100-200
μm. Fibers with simple to minutely
bordered pits (libriform fibers). All ray
cells upright and square.

›T ›T

Tangential section
Rays uniseriate and up to 10 seriate.

Wood basic density: 0,55 g/cm³

›T ›T

145
Hirtellina lobelii (DC.) Dittrich = Staehelina lobelii DC.
Asteraceae

PLANT DESCRIPTION
Subshrub up to 60 cm high, with nu-
merous erect, slender stems, branched
only in the region of inflorescences. A
rare indigenous species on Cyprus, oc-
curring on fissures of limestone rocks
(500-800 m alt.). Found also in south-
ern Turkey, Lebanon and Crete.

TWIG ANATOMY
Bark
No slide available.

Xylem
As stem wood.
Hirtellina lobelii (DC.) Dittrich

Pith
Pith shape round. Heterogeneous pith.
Thick-walled parenchyma cells present.
Cells dimorphic. Pits in transverse and
in longitudinal cell walls. Vascular bun-
dles clearly separate. Tracheary ele-
ments of metaxylem in distinct radial
rows.

›T ›T

146
Stem xylem: 1 4 7 9.1 11 13 21 22 25 40.2 45 50.1 52.3 61 69 70.3 78 79 89 98 109 114 136
Twig bark: No slide available.
Pith: P1 P3 P3.3 P4 P9 P9.1 P10.1 P12

Asteraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted latewood fibers. Wood
semi-ring-porous. Vessels in diagonal
and radial pattern. Vessels in radial
multiples of 2 to 4 elements and in
clusters. Mean tangential diameter of
earlywood vessels 20-50 μm, 100-200
vessels per mm2. Vessels in two dis-
tinct diameter classes. Fibers thin- to
thick-walled. Radial flat marginal fib-
ers. Axial parenchyma scanty paratra-
cheal to vasicentric. Apotracheal pa-
renchyma in marginal or in seemingly
marginal bands. Less than 4 rays per
›T ›T
mm.

Radial section
Simple perforation plates. Inter-vessel
pits opposite and alternate, small (4-7
μm in diameter). Earlywood vessel
element length 100-200 μm. Fibers
with simple to minutely bordered pits
(libriform fibers). Rays with procum-
bent, square and upright cells mixed
throughout the ray. Prismatic crystals
present.

›T ›T

Tangential section Hirtellina lobelii (DC.) Dittrich


Rays commonly 4 to 10 seriate.

Wood basic density: 0,69 g/cm³

›T ›T

147
Limbarda crithmoides (L.) Dumort. subsp. longifolia
(Arcang.) Greuter = Inula crithmoides L.
Asteraceae

PLANT DESCRIPTION
Subshrub up to 80 cm high. Leaves
numerous, linear or oblanceolate, suc-
culent. Distributed in throughout west-
ern Europe, the Mediterranean coasts
and the Atlantic islands. Indigenous to
Cyprus, found near salt lakes and salt
marshes in costal areas (0-50 m alt.).
Limbarda crithmoides (L.) Dumort. subsp. longifolia (Arcang.) Greuter

TWIG ANATOMY
Bark
Groups of sieve tube present. Some
rays become dilated. Fibers grouped.
Sclereids scattered or irregularly dis-
persed. Acicular crystals present. Dark
staining substances in the transition
zone between cortex and phellem.
Phellem homogeneous, distinct in po-
larized light.

Xylem
Vessels in radial multiples of 4 or
more. Mean tangential diameter of ear-
lywood vessels less than 20 μm. Less
than 4 rays per mm. Rays uniseriate
and up to 3 cells wide. ›T

Pith
Pith shape round. Heterogeneous pith.
Unlignified cells present. Crystal sand
present. Pits in transverse and in lon-
gitudinal cell walls. Vascular bundles
clearly separate. Axial cells in regular
rows (radial section).

›T ›T

148
Stem xylem: 2.1 5 8 9 9.1 13 21 22 25 30 40.2 45 50.2 52.3 60 61 69 78 96 99 102 103 105 115
Twig bark: B1 B6 B9 B14 B15 B18 B21 B31 B14.3
Pith: P1 P3 P3.4 P6.4 P9 P9.1 P10.1 P13

Asteraceae
STEM ANATOMY
Transverse section
Only one ring in the observed sample.
Wood diffuse-porous. Vessels pre-
dominantly solitary or in short radial
multiples, showing a dendritic pattern.
Mean tangential diameter of earlywood
vessels 20-50 μm. Vessels of two dis-
tinct diameter classes. More than 200
vessels per mm2. Fibers thin- to thick-
walled. Axial parenchyma scanty para-
tracheal. Rays 4-12 per mm.

Limbarda crithmoides (L.) Dumort. subsp. longifolia (Arcang.) Greuter


›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits opposite and alternate, small (4-7
μm in diameter). Vessel-ray pits with
distinct borders, similar to interves-
sel pits in size and shape throughout
the ray cell. Earlywood vessel element
length 100-200 μm. Vascular and/or
vasicentric tracheids present. Fibers
with simple to minutely bordered pits
(libriform fibers). All ray cells upright
and square.

›T ›T

Tangential section
Rays of two distinct sizes: uniseriate
and more than 10 seriate, the latter
more than 1 mm high.

Wood basic density: 0,47 g/cm³

›T ›T

149
Phagnalon rupestre (L.) DC. subsp. rupestre
African Fleabane
Asteraceae

PLANT DESCRIPTION
Erected or sprawling, much-branched
subshrub up to 50 cm high. Young
stems densely white-tomentose, be-
coming glabrous and dark brown on
old growths. Indigenous to eastern
Mediterranean region. In Cyprus it oc-
curs in garigue, on dry and rocky sites
(0-900 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube present. Some
rays become dilated. Sclerenchyma
cells in phloem and in cortex. Fibers
in tangentially elongated groups with
sieve tube and parenchyma cell.

Xylem
Larger rays commonly 4 to 10 seriate.
Phagnalon rupestre (L.) DC. subsp. rupestre

›T

Pith
Pith shape roundh. Cells dimorphic.
Heterogeneous pith. Unlignified cells
present. Cell content present (dark
staining substances). Pits in transverse
and in longitudinal cell walls. Vascular
bundles clearly separate. Axial cells in
regular rows (radial section).

›T ›T

150
Stem xylem: 1 4 5 9 13 22 25 30 40.2 45 50.2 52.3 58 60 61 69 70 78 79 89 96 98 102 103 105 115
Twig bark: B1 B6 B7 B14
Pith: P1 P3 P3.4 P4.1 P5 P9 P9.1 P10.1 P13

Asteraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted latewood fibers and by
marginal parenchyma bands. Wood
semi-ring-porous to diffuse-porous.
Vessels predominantly solitary. Mean
tangential diameter of earlywood ves-
sels 20-50 μm. More than 200 vessels
per mm2. Vessels of two distinct diam-
eter classes. Gums and other deposits
in heartwood vessels. Fibers thick- to
very thick-walled. Axial parenchyma
scanty paratracheal to vasicentric.
Apotracheal parenchyma in seemingly
›T ›T
marginal bands. Rays 4-12 per mm.

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 100-
200 μm. Vascular and/or vasicentric
tracheids present. Fibers with simple

Phagnalon rupestre (L.) DC. subsp. ruperstre


to minutely bordered pits (libriform fib-
ers). All ray cells upright and square.

›T ›T

Tangential section
Rays of two distinct sizes: uniseri-
ate and up to 10 seriate. Larger rays
height commonly more than 1 mm.

Wood basic density: 0,90 g/cm³

›T ›T

151
Ptilostemon chamaepeuce (L.) Less. subsp. cyprius (Greuter) B. Slavík
et Chrtek = Ptilostemon chamaepeuce (L.) Less. var. cyprius Greuter
Asteraceae

PLANT DESCRIPTION
Evergreen shrub up to 1,3 m high.
Young branches densely white-tomen-
tose, old branches greyish. The sub-
species is endemic to Cyprus, occur-
ring on rocky slopes and rock fissures
on calcareous and indigenous forma-
tions (50-1600 m alt.).
Ptilostemon chamaepeuce (L.) Less. subsp. cyprius (Greuter) B. Slavík et Chrtek

TWIG ANATOMY
Bark
Groups of sieve tube in radial rows.
Collapsed sieve tubes present. Fibers
in tangential rows, scattered or irregu-
larly dispersed. Cortex consist of large
and small thin-walled parenchyma
cells.

Xylem
Earlywood vessels length 100-200 μm.
All ray cells upright and square.

›T

Pith
Pith shape round. Cells dimorphic. Cell
content present (dark staining sub-
stances). Crystal druses present. Pits
in transverse and in longitudinal cell
walls. Vascular bundles clearly sepa-
rate. Tracheary elements of metaxylem
in distinct radial rows. Axial cells in
regular rows (radial section).

›T ›T

152
Stem xylem: 1 4 5 7 8 11 13 22 26 30 40.2 45 50.2 53.1 56 58 61 70 70.3 78 79 89 96 97 103 109 115
Twig bark: B3 B4 B9 B12 B5.3
Pith: P1 P4.1 P5 P6.2 P9 P9.1 P10.1 P12 P13

Asteraceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by radi-
ally flatted latewood fibers and by initial
marginal parenchyma bands. Wood
semi-ring-porous to diffuse-porous.
Vessels in radial/diagonal pattern, pre-
dominantly in clusters. Mean tangential

Ptilostemon chamaepeuce (L.) Less. subsp. cyprius (Greuter) B. Slavík et Chrtek


diameter of earlywood vessel lumina
20-50 μm. More than 200 vessels per
mm2. Vessels of two distinct diameter
classes. Tyloses and deposits in heart-
wood vessels. Fibers very thick-walled.
Axial parenchyma scanty paratracheal
to vasicentric. Rays 4-12 per mm.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate. Intervessel pits me-
dium (7-10 μm in diameter). Vessel-
ray pits with distinct borders, similar
to intervessel pits in size and shape
throughout the ray cell. Earlywood ves-
sel element length 200-500 μm. Fibers
with simple to minutely bordered pits
(libriform fibers). Rays with procum-
bent, square and upright cells mixed
throughout the ray.

›T ›T

Tangential section
Rays of two distinct sizes: uniseriate
and 2-3 seriate. Larger rays more than
1 mm high.

Wood basic density: 0,69 g/cm³

›T ›T

153
Berberis cretica L.
Barberry
Berberidaceae

PLANT DESCRIPTION
Deciduous shrub, up to 2 m high. Bark
reddish-brown. Leaves of long shoots
transformed into three-partite spines.
Leaves in fascicles in the axils of the
spines. Indigenous to Cyprus, growing
in pine forests and on screes (900-
1950 m alt.). It also occurs in Greece,
Crete, the Aegean islands and Turkey.

TWIG ANATOMY
Bark
Groups of sieve tube in tangential
rows. Only some rays become dilated.
Sclereids in tangential rows and scat-
tered or irregularly dispersed. Single
lignified cells scattered in phellem.

Xylem
Mean tangential diameter of earlywood
vessels 20-50 μm. Larger rays com-
monly 4 to 10 seriate. Rays 4-12 per
mm.

›T

Pith
Pith shape round. Heterogeneous
pith. Thick-walled cells present. Pits
in transverse and in longitudinal cell
walls. Vascular bundles clearly sepa-
rate. Axial cells arranged almost in
regular rows (radial section). ›T
Berberis cretica L.

›T

154
Stem xylem: 1 3 7 8 9.1 11 13 22 25 30 31 36 40.2 48 52.3 60 61 69 78 99 102 109 114
Twig bark: B1 B2 B6 B15 B17 B18 B34
Pith: P1 P3 P3.4 P9 P9.1 P10.1 P13

Berberidaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
the difference in vessel size between
latewood and earlywood. Wood ring-
porous. Vessels in diagonal and radial
pattern. Vessels in radial multiples of
2 to 4 and in clusters. Mean tangential
diameter of earlywood vessels 20-50
μm, 20-40 vessels per mm2. Fibers
thin- to thick-walled. Axial parenchyma
scanty paratracheal. Less than 4 rays
per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in diame-
ter). Vessel-ray pits with distinct bor-
ders, similar to intervessel pits in size
and shape throughout the ray cell, or
rounded with large apertures. Helical
thickenings mainly in narrower vessel
elements. Earlywood vessel element
length 100-200 μm. Vascular and/or
vasicentric tracheids present. Fibers
with simple to minutely bordered pits
(libriform fibers). Rays with procum-
bent, square and upright cells mixed
throughout the ray.

›T ›T

Tangential section
Larger rays commonly more than 10
seriate. Ray height more than 1 mm.
Berberis cretica L.

Wood basic density: 0,69 g/cm³

›T ›T

155
Alnus orientalis Decne.
Oriental Alder
Betulaceae

PLANT DESCRIPTION
Deciduous tree up to 20 m high. Twigs
rather slender, glabrous. Leaves al-
ternate, ovate or oblong, with 2-3 cm
long petiole. Indigenous to Cyprus,
found abundantly along rivers and
main streams with considerable water
flow (0-1550 m alt.). Also indigenous
to Turkey, Syria and Lebanon.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tube present. Fibers in tangential
rows, sclereids scattered or irregularly
dispersed. Prismatic crystals and crys-
tal druses present. Cortex homogene-
ous, distinct in polarized light.

Xylem
Vessels in short radial multiples.
Solitary vessels outline angular. Fibers
thin-walled.

›T

Pith
Pith shape triangular. Cells dimorphic.
Prismatic crystals present. Pits grouped
in transverse and in longitudinal cell
walls. Vascular bundles clearly sepa-
Alnus orientalis Decne.

rate. Tracheary elements of metaxylem


in distinct radial rows. Axial cells in
regular rows (radial section).

›T ›T

156
Stem xylem: 1 5 9.1 10 11 14 16 21 24 30 41 50.1 53.1 61 69 70.2 76 96 101 104 116.1
Twig bark: B1 B4 B12 B18 B20 B22 B31 B14.3
Pith: P1.2 P4 P6 P9 P9.1 P9.2 P10.1 P12 P13

Betulaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood diffuse-porous. Vessels in
radial multiples of 2 to 4 or more ele-
ments, and in clusters. Mean tangential
diameter of earlywood vessel lumina
50-100 μm, 100-200 vessels per mm2.
Vessel-ray pits with distinct borders,
similar to intervessel pits in size and
shape throughout the ray cell. Fibers
thin- to thick-walled. Tension wood
present. Axial parenchyma diffuse.
Rays 12-20 per mm.
›T ›T

Radial section
Scalariform perforation plates with
more than 10 bars. Inter-vessel pits
opposite, minute (less than 4 μm in di-
ameter). Earlywood vessels length 200-
500 μm. Fibers with simple to minutely
bordered pits (libriform fibers). All ray
cells procumbent.

›T ›T

Tangential section
Rays exclusively uniseriate. Aggregate
rays present.
Alnus orientalis Decne.

Wood basic density: 0,36 g/cm³

›T ›T

157
Corylus avellana L.
European Hazel, Common Hazel
Betulaceae

PLANT DESCRIPTION
Deciduous shrub up to 15 m high.
Leaves are alternate, rounded, 6–12
cm long and across, softly hairy on
both surfaces. Fruit a one-seeded,
ovoid nut. Naturalized in Cyprus, it
occurs along streams and on cultivated
ground (500-1200 m alt.). Native to
Europe and western Asia.

TWIG ANATOMY
Bark
Collapsed sieve tubes present. Distinct
ray dilatation. Sclerenchyma cells in
phloem and in cortex. Fibers and scle-
reids grouped. Prismatic crystals and
crystal druses present. Phellem homo-
geneous, distinct in polarized light.

Xylem
As stem wood.

›T

Pith
Pith shape round. Medullary sheath
consist of thick-walled cells. Crystal
druses present. Pits in transverse and
in longitudinal cell walls. Vascular bun-
dles clearly separate. Tracheary ele-
ments of metaxylem in distinct radial
Corylus avellana L.

rows.

›T ›T

158
Stem xylem: 1 4 5 9 9.1 10 14 15 22 25 30 36 40.2 50.1 53.2 60 61 69 70.2 76 86 97 101 104 106 116.1
Twig bark: B4 B5 B7 B9 B14 B15 B19 B20 B22 B31 B14.3
Pith: P1 P2 P3 P6.2 P9 P9.1 P10.1 P12

Betulaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers and by a lack of vessels in termi-
nal latewood. Wood semi-ring-porous
to diffuse-porous. Vessels solitary and
in radial multiples up to 4 or more.
Mean tangential diameter of earlywood
vessels 20-50 μm. Vessels per mm2
100-200. Fibers thin- to thick-walled.
Tension wood present. Apotracheal pa-
renchyma diffuse and in narrow bands
or lines up to three cells wide. Rays
12-20 per mm.
›T ›T

Radial section
Scalariform perforation plates with
less than 10 bars. Inter-vessel pits
alternate, small (4-7 μm in diameter).
Vessel-ray pits with distinct borders,
similar to intervessel pits in size and
shape throughout the ray cell. Helical
thickenings in vessel elements present.
Earlywood vessel element length more
than 500 μm. Vascular and/or vasi-
centric tracheids present. Fibers with
simple to minutely bordered pits (libri-
form fibers). Some rays with procum-
bent cells, larger rays with body cells
procumbent and one row of square
marginal cells.
›T ›T

Tangential section
Ray width predominantly 1 to 3 cells.
Aggregate rays present.
Corylus avellana L.

Wood basic density: 0,47 g/cm³

›T ›T

159
Ostrya carpinifolia L.
Hop Hornbeam
Betulaceae

PLANT DESCRIPTION
A deciduous tree up to 24 m high
with a conical or irregular crown and a
scaly, rough bark. Leaves alternate and
double-toothed, 3–10 cm long. Fruit in
pendulous clusters with 6–20 seeds. It
occurs in central and souther Europe
(300-1500 m alt.).

TWIG ANATOMY
Bark
Collapsed sieve tubes present. Fibers
and sclereids in groups. Prismatic crys-
tals and crystal druses present. Phloem
uniform. Cortex homogeneous.

Xylem
As stem wood.

›T

Pith
Pith shape round. Medullary sheath
present. All cells thick-walled. Cells di-
morphic. Pits in transverse and in lon-
gitudinal cell walls. Vascular bundles
clearly separate. Tracheary elements of
Ostrya carpinifolia L.

metaxylem in distinct radial rows.

›T ›T

160
Stem xylem: 1 5 9 10 13 22 30 36 40.2 49 53.1 60 61 69 76 77 97 104 116.2 136 136.1
Twig bark: B4 B9 B14 B15 B19 B20 B22 B8 B14.1
Pith: P1 P2 P3.3 P4 P9 P9.1 P10.1 P12

Betulaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted latewood fibers and
by marginal parenchyma cells. Wood
diffuse-porous. Vessels predominantly
solitary. Vessels in radial multiples of 4
or more elements common. Mean tan-
gential diameter of earlywood vessels
20-50 μm, 40-100 vessels per mm2.
Fibers thin- to thick-walled. Axial pa-
renchyma diffuse in aggregates. More
than 20 rays per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate. Vessel-ray pits with
distinct borders, similar to intervessel
pits in size and shape throughout the
ray cell. Helical thickenings in vessel
elements present. Earlywood vessels
length 200-500 μm. Vascular and/or
vasicentric tracheids present. Fibers
with simple to minutely bordered pits
(libriform fibers). All ray cells procum-
bent. Prismatic crystals in ray paren-
chyma cells.

›T ›T

Tangential section
Ray width predominantly 1 to 3 cells.
Ostrya carpinifolia L.

Wood basic density: 0,82 g/cm³

›T ›T

161
Echium angustifolium Mill.
Narrow–leaved Bugloss, Hispid Viper’s-bugloss
Boraginaceae

PLANT DESCRIPTION
Erect or sprawling perennial subshrub
10-60 cm high. An Eastern Mediterra-
nean species, distributed From Greece
to Egypt and Libya. Indigenous to
Cyprus, where it is locally common on
rocky and sandy seashores, roadsides,
dry banks and hillsides (0-1000 m alt.).

TWIG ANATOMY
Bark
Phloem uniform. Cell content in pa-
renchyma cells of the cortex. Phellem
consist of large, collapsed, thin-walled
cells.

Xylem
Only one ring in the observed sample.
Rays absent or extremely rare or not to
recognizable.

›T

Pith
Pith not visible in polarized light. Pith
Echium angustifolium Mill.

shape round. Unlignified cells pre-


sent. Vascular bundles not distinct.
Tracheary elements of metaxylem in
distinct radial rows.

›T ›T

162
Stem xylem: 1 5 9 11 13 22 25 30 40.2 45 49 52.3 58 62 69 78 89 89.1 98 100.2 109 115
Twig bark: B8 B28
Pith: P1 P3.4
.2 P12

Boraginaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
initial marginal parenchyma bands.
Wood diffuse-porous. Vessels solitary
and small clusters. Mean tangential
diameter of earlywood vessels 20-50
μm, 40-100 vessels per mm2. Ves-
sels of two distinct diameter classes.
Gums and other deposits in heartwood
vessels. Fibers thin- to thick-walled.
Axial parenchyma scanty paratracheal.
Apotracheal parenchyma cells in mar-
ginal or in seemingly marginal bands,
thin-walled, dark in polarized light.
Rays 4-12 per mm.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 100-
200 μm. Fibers with distinctly bor-
dered pits (fiber tracheids). Rays with
procumbent, square and upright cells
mixed throughout the ray.

›T ›T

Tangential section
Larger rays commonly 4 to 10 seriate.
Echium angustifolium Mill.
Some rays partially disappear in polar-
ized light.

Wood basic density: 0,41 g/cm³

›T ›T

163
Lithodora hispidula (Sm.) Griseb. subsp. versicolor Meikle
Boraginaceae

PLANT DESCRIPTION
Evergreen, much branched shrub,
1-1,5 m high, with hairy shoots. The
flowering shoots are spinescent when
dead. Leaves with long and stiff hairs.
Indigenous to Cyprus, very common
in many areas with garigue vegetation
or open forests (0-1000 m alt.). Indig-
enous also in Turkey and Syria.

TWIG ANATOMY
Lithodora hispidula (Sm.) Griseb. subsp. versicolor Meikle

Bark
Groups of sieve tube present. Some
rays become dilated. Sclerenchyma
cells in phloem and in cortex. Fibers
and sclereids scattered or irregularly
dispersed. Sclereids in tangential rows,
too. Cell content in parenchyma cells.

Xylem
Vessels predominately solitary. Mean
tangential diameter of earlywood
vessels 20-50 μm. Fibers very thick-
walled.

›T

Pith
Pith shape round. Heterogeneous
pith. Thick-walled parenchyma cells
present. Cells dimorphic. Cell content
present (dark staining substances). Pits
in transverse and in longitudinal cell
walls. Vascular bundles clearly sepa- ›T
rate to not distinct. Tracheary elements
of metaxylem in distinct radial rows.

›T

164
Stem xylem: 1 4 5 9.1 11 13 21 22 25 30 36 39.1 40.1 45 50.1 52.3 62 69 76 78 89 98 109 114
Twig bark: B1 B6 B7 B9 B13 B15 B17 B18 B28
Pith: P1 P3 P3.3 P4 P5 P9 P9.1 P10.1 P10.2 P12

Boraginaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands and by a
lack of vessels in terminal latewood.
Wood semi-ring-porous to diffuse-
porous. Vessels in short radial multi-
ples and in clusters. Vessels cell wall
thick (more than 2 μm). Vessels of
two distinct diameter classes. Mean
tangential diameter of earlywood ves-
sels less than 20 μm, 100-200 vessels
per mm2. Fibers thin- to thick-walled.
Axial parenchyma diffuse and scanty
paratracheal. Apotracheal parenchyma
›T ›T
in marginal or in seemingly marginal
bands. Less than 4 rays per mm.

Radial section
Simple perforation plates. Inter-vessel

Lithodora hispidula (Sm.) Griseb. subsp. versicolor Meikle


pits opposite and alternate, small (4-7
μm in diameter). Vessel-ray pits with
distinct borders, similar to intervessel
pits in size and shape throughout the
ray cell. Helical thickenings in vessel
elements present. Earlywood vessel
element length 100-200 μm. Fibers
with distinctly bordered pits (fiber tra-
cheids). Rays with procumbent, square
and upright cells mixed throughout the
ray.

›T ›T

Tangential section
Larger rays commonly 4 to 10 seriate.

Wood basic density: 0,65 g/cm³

›T ›T

165
Onosma caespitosa Kotschy
Boraginaceae

PLANT DESCRIPTION
Loosely tufted perennial subshrub up
to 30 cm high; twigs covered with rigid
hairs. Endemic to Cyprus, growing on
cliffs of limestone along the Pentadak-
tylos range (300-900 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclereids scattered
or irregularly dispersed. Cortex consist
of large and small thin-walled paren-
chyma cells. Prismatic crystals present.
Lignified cells in cortex.

Xylem
As stem wood.

›T

Pith
Onosma caespitosa Kotschy

Pith shape round. Medullary sheath


present. Unlignified cells present.
Laticifers or intercellular canals pre-
sent. Vascular bundles not distinct.

›T ›T

166
Stem xylem: 2 5 7 9 11 13 21 24 36 40.1 45 49 52.3 69 79.1 117
Twig bark: B1 B4 B18 B20 B34
Pith: P1 P2 P3.4 P7 P10.2

Boraginaceae
STEM ANATOMY
Transverse section
Growth ring indistinct or absent. Wood
diffuse-porous. Vessels in diagonal and
radial pattern. Vessels solitary or in
clusters. Mean tangential diameter of
earlywood vessels less than 20 μm, 40-
100 vessels per mm2. Vessels of two
distinct diameter classes. Fibers thin-
to thick-walled. Parenchyma pervasive.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits opposite, minute (less than 4 μm
in diameter). Helical thickenings in ves-
sel elements present. Earlywood vessel
element length 100-200 μm.

Tangential section Onosma caespitosa Kotschy


Wood rayless.

Wood basic density: 0,50 g/cm³

›T

167
Onosma fruticosa Sm.
Cyprus Golden-Drop
Boraginaceae

PLANT DESCRIPTION
Much branched, evergreen shrub 20-
80 cm high, with stiffly hairy shoots
and leaves. Endemic to Cyprus, found
in many areas with garigue vegetation
(0-900 m alt.).

TWIG ANATOMY
Bark
Phloem uniform. Dark staining sub-
stances in parenchyma cells of the
cortex. Cortex uniform.

Xylem
As stem wood. The first growth ring
distinctly differs to the others because
of a lack in vessels and abundance of
thick-walled fibers.

›T

Pith
Pith shape round. Thick-walled paren-
chyma cells present. Cell content pre-
sent (dark staining substances). Pits in
transverse cell walls. Vascular bundles
Onosma fruticosa Sm.

clearly separate. Tracheary elements of


metaxylem in distinct radial rows.

›T ›T

168
Stem xylem: 1 4 7 11 13 22 24 40.2 45 50.1 52.3 58 62 69 89 89.1 117
Twig bark: B8 B28 B14.1
Pith: P1 P3.3 P5 P9.1 P10.1 P12

Boraginaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands. Wood
semi-ring-porous. Vessels in diagonal
and radial pattern, predominantly in
clusters. Mean tangential diameter of
earlywood vessels 20-50 μm, 100-200
vessels per mm2. Vessels of two dis-
tinct diameter classes. Gums and other
deposits in heartwood vessels. Fib-
ers thin- to thick-walled. Apotracheal
parenchyma in marginal or in seem-
ingly marginal bands, dark in polarized
light.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, minute (less than 4 μm
in diameter). Earlywood vessel element
length 100-200 μm. Fibers with dis-
tinctly bordered pits (fiber tracheids).

›T ›T

Tangential section
Wood rayless.
Onosma fruticosa Sm.

Wood basic density: 0,61 g/cm³

›T ›T

169
Onosma mitis Boiss. et Heldr.
Boraginaceae

PLANT DESCRIPTION
Subshrub, 15-50 cm high; stems and
leaves densely hairy. Native of Cyprus
and southern Turkey, rather unusual,
confined to the southwestern part of
the Troodos range (400-900 m alt.).

TWIG ANATOMY
Bark
Phloem uniform. Phellem consist of
squared cells and collapsed cells.

Xylem
As stem wood.

›T

Pith
Onosma mitis Boiss. et Heldr.

Pith not visible in polarized light. Pith


shape round. Heterogeneous pith.
Unlignified cells present. Vascular bun-
dles not distinct.

›T

›T

170
Stem xylem: 1 5 9 9.1 11 13 22 25 40.1 40.2 45 50.1 52.3 58 61 69 78 89 89.1 117
Twig bark: B8 B32
Pith: P0.1 P1 P3 P3.4 P10.2

Boraginaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands. Wood
diffuse-porous. Vessels solitary and
in small groups. Mean tangential di-
ameter of earlywood vessels 20-50
μm, 100-200 vessels per mm2. Ves-
sels of two distinct diameter classes.
Gums and other deposits in heartwood
vessels. Fibers thin- to thick-walled.
Axial parenchyma scanty paratracheal.
Apotracheal parenchyma in marginal
or in seemingly marginal bands. Paren-
chyma marginal thin-walled, dark in
polarized light.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in diam-
eter). Earlywood vessel element length
100-200 μm. Fibers with simple to mi-
nutely bordered pits (libriform fibers).

›T ›T

Tangential section Onosma mitis Boiss. et Heldr.


Wood rayless.

Wood basic density: 0,47 g/cm³

›T ›T

171
Alyssum akamasicum B.L. Burtt
Akamas Alison
Brassicaeae

PLANT DESCRIPTION
A subshrub or perennial herb with
erect or decumbent stems up to 40
cm long. Stems and leaves densely
covered with stellate hairs. Endemic to
Cyprus. It occurs on dry, rocky hill-
sides and shrubberies with serpentine
rocks, in the Akamas peninsula (0-400
m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclereids in radial
rows, scattered or irregularly dis-
persed, sometimes in groups. Cortex
consist of large parenchyma cells,
distinct in polarized light.

Xylem
As stem wood.
Alyssum akamasicum B.L. Burtt

Pith
No slide available.

›T

172
Stem xylem: 1 5 9 9.1 13 21 22 25 39.1 40.1 50.2 52.2 61 69 70 78 89 117
Twig bark: B1 B4 B16 B18 B19 B31 B33
Pith: No slide available.

Brassicaeae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands. Wood
diffuse-porous. Vessels solitary and
in radial multiples of 2 to 4. Vessels
cell wall thick (more than 2 μm). Mean
tangential diameter of earlywood ves-
sels less than 20 μm. More than 200
vessels per mm2. Fibers thick- to very
thick-walled. Axial parenchyma scanty
paratracheal, and in marginal or seem-
ingly marginal bands.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits opposite and alternate, small (4-7
μm in diameter). Earlywood vessel ele-
ment length 50-100 μm. Fibers with
simple to minutely bordered pits (libri-
form fibers).

›T ›T

Tangential section Alyssum akamasicum B.L. Burtt


Wood rayless.

Wood basic density: 0,43 g/cm³

›T ›T

173
Alyssum chondrogynum B.L. Burtt
Kakomallis Alison
Brassicaeae

PLANT DESCRIPTION
A robust subshrub up to 40 cm high.
Stem and leaves with dense, stellate
hairs. Endemic to Cyprus. It is found
on rocky mountainsides with serpen-
tine ultrabasic rocks (300-100 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Some rays become
dilated. Sclerenchyma cells in phloem
and in cortex. Fibers in radial groups.
Cortex consist of large and small thin-
walled parenchyma cells. Phellem ho-
mogeneous, distinct in polarized light.

Xylem
As stem wood.
Alyssum chondrogynum B.L. Burtt

Pith
No slide available.

›T

174
Stem xylem: 1 5 9 9.1 13 22 24 31 39.1 40.1 50.1 52.2 61 70 78 89.1 98 100.2 109 115
Twig bark: B1 B4 B6 B7 B9 B11 B31 B33
Pith: No slide available.

Brassicaeae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands. Wood
diffuse-porous. Vessels predominantly
solitary or in short radial multiples.
Vessels cell wall thick (more than 2
μm). Mean tangential diameter of
earlywood vessels less than 20 μm,
100-200 vessels per mm2. Fibers very
thick-walled. Axial parenchyma scanty
paratracheal, and in marginal or seem-
ingly marginal bands. Rays 4-12 per
mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, minute (less than 4 μm
in diameter). Vessel-ray pits rounded
or angular with large apertures. Early-
wood vessels length 50-100 μm. Fibers
with simple to minutely bordered pits
(libriform fibers). Rays with square and
upright cells mixed throughout the ray.

Alyssum chondrogynum B.L. Burtt


›T ›T

Tangential section
Larger rays commonly 4 to 10 seriate.
Rays disappear in polarized light.

Wood basic density: 0,68 g/cm³

›T

175
Alyssum cypricum Nyar.
Cyprus Alison
Brassicaeae

PLANT DESCRIPTION
A subshrub up to 30 cm high. Stems
and leaves densely covered with stel-
late hairs. A species native to Cyprus.
It grows on rocky mountainsides of
serpentinised, ultrabasic rocks, on
the central part of the Troodos range
(1100-1950 m alt.). It is indigenous
also to Turkey.

TWIG ANATOMY
Bark
Collapsed sieve tubes present. Fibers
in large, radially arranged groups. Lo-
cally groups of unlignified parenchyma
cells.

Xylem
As stem wood.

Pith
No slide available.
Alyssum cypricum Nyar.

›T

176
Stem xylem: 1 5 9.1 11 13 22 25 39.1 40.1 50.1 52.2 58 61 70 78 97 105 114 133.2
Twig bark: B4 B11
Pith: No slide available.

Brassicaeae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands. Wood
diffuse-porous. Vessels in radial mul-
tiples of 2 to 4 or in clusters. Vessels
cell wall thick (more than 2 μm). Mean
tangential diameter of earlywood ves-
sels less than 20 μm, 100-200 vessels
per mm2. Gums and other deposits in
heartwood vessels. Fibers very thick-
walled. Axial parenchyma scanty para-
tracheal. Less than 4 rays per mm. In-
cluded phloem: concentric continuous.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in diam-
eter). Earlywood vessel element length
50-100 μm. Fibers with simple to mi-
nutely bordered pits (libriform fibers).
All ray cells upright.

›T ›T

Tangential section
Ray width predominantly 1 to 3 cells.
Alyssum cypricum Nyar.

Wood basic density: 0,65 g/cm³

›T ›T

177
Alyssum troodi Boiss.
Troodos Alison
Brassicaeae

PLANT DESCRIPTION
A subshrub up to 30 cm high. Stems
and leaves with densely covered with
stellate hairs. Endemic to Cyprus.
It grows on rocky mountainsides of
serpentinised, ultrabasic rocks, around
Chionistra in the Troodos range (1200-
1950 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclereids in radial
rows, scattered or irregularly dispersed
or in groups. Cortex consist of large
parenchyma cells. Phellem homogene-
ous, distinct in polarized light.

Xylem
As stem wood.

›T

Pith
Pith not visible in polarized light. Pith
shape triangular to roundish. Cells di-
morphic. Pits in transverse and in lon-
gitudinal cell walls. Pits of two distinct
Alyssum troodi Boiss.

size. Vascular bundles in the pith.

›T

178
Stem xylem: 1 5 9 9.1 13 20 22 25 39.1 40.1 50.2 52.2 61 69 70 78 89.1 89.2 117
Twig bark: B1 B4 B16 B18 B19 B31 B33
Pith: P1.1 P1.2 P4 P9 P9.1 P9.3 P10

Brassicaeae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands. Wood
diffuse-porous. Vessels predominantly
solitary or in radial multiples of 2 to
4. Vessels cell wall thick (more than 2
μm). Mean tangential diameter of ear-
lywood vessels less than 20 μm, more
than 200 vessels per mm2. Fibers thick-
to very thick-walled. Axial parenchyma
scanty paratracheal. Parenchyma mar-
ginal thin-walled, dark in polarized
light. Ring shake common.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits scalariform and alternate, small
(4-7 μm in diameter). Earlywood vessel
element length 50-100 μm. Fibers with
simple to minutely bordered pits (libri-
form fibers).

›T ›T

Tangential section
Wood rayless.
Alyssum troodi Boiss.

Wood basic density: 0,56 g/cm³

›T ›T

179
Arabis cypria Holmboe
Cyprus Rock-Cress
Brassicaeae

PLANT DESCRIPTION
A subshrub up to 30 cm high. Leaves
with long, stellate hairs. Endemic to
Cyprus, growing in crevices of lime-
stone cliffs and rocks (300-900 m alt.),
mainly on limestone along the Pen-
tadaktylos mountain range.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes arranged in tangential
groups. Cortex consist of large and
small thin-walled parenchyma cells.
Phellem homogeneous, distinct in po-
larized light.

Xylem
As stem wood.

Pith
No slide available.
Arabis cypria Holmboe

›T

180
Stem xylem: 1 5 9 9.1 11 13 20 22 25 39.1 40.1 50.2 53.1 61 69 70 79 89.1 89.2 117
Twig bark: B1 B4 B31 B33
Pith: No slide available.

Brassicaeae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands. Wood
diffuse-porous. Vessels solitary, in ra-
dial multiples of 2 to 4 on in clusters.
Vessels cell wall thick (more than 2
μm). Mean tangential diameter of ear-
lywood vessels less than 20 μm. More
than 200 vessels per mm2. Fibers thick-
to very thick-walled. Axial parenchyma
vasicentric. Parenchyma marginal thin-
walled, dark in polarized light. Ring
shake. Included phloem: concentric
continuous.
›T

Radial section
Simple perforation plates. Inter-vessel
pits scalariform and alternate, small
(4-7 μm in diameter). Earlywood ves-
sel element length 200-500 μm. Fibers
with simple to minutely bordered pits
(libriform fibers).

Tangential section
Wood rayless.
Arabis cypria Holmboe

Wood basic density: 0,40 g/cm³

›T

181
Arabis purpurea Sm.
Purple Rock-Cress
Brassicaeae

PLANT DESCRIPTION
A subshrub up to 30 cm high. Leaves
with short stellate hairs. Locally very
common Cypriot endemic. It occurs
in crevices of igneous and calcareous
rocks (400-1800 m alt.), mainly along
the Troodos range.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclereids in
groups. Phellem distinct in polarized
light. Lignified cells in phellem.

Xylem
As stem wood.

›T

Pith
Pith not visible in polarized light. Pith
shape polygonal. Sclereids scattered or
irregularly dispersed. Cells dimorphic.
Pits in transverse and in longitudinal
cell walls. Pits of two distinct size.
Arabis purpurea Sm.

Vascular bundles at the periphery of


the pith.

›T ›T

182
Stem xylem: 1 5 9 11 13 20 22 25 39.1 40.1 50.2 53.1 61 69 78 89.1 89.2 117 133.2
Twig bark: B1 B4 B19 B31 B33 B34
Pith: P1.1 P3.1 P4 P9 P9.1 P9.3 P10

Brassicaeae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands. Wood
diffuse-porous. Vessels solitary and in
clusters. Vessels cell wall thick (more
than 2 μm). Mean tangential diameter
of earlywood vessels less than 20 μm.
More than 200 vessels per mm2. Fibers
thin- to thick-walled. Axial parenchyma
scanty paratracheal. Parenchyma mar-
ginal thin-walled, dark in polarized
light. Ring shake. Included phloem:
concentric continuous.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits scalariform. Inter-vessel pits al-
ternate, small (4-7 μm in diameter).
Earlywood vessel element length 200-
500 μm. Fibers with simple to minutely
bordered pits (libriform fibers).

›T ›T

Tangential section
Wood rayless.
Arabis purpurea Sm.

Wood basic density: 0,33 g/cm³

›T ›T

183
Brassica hilarionis Post
Brassicaeae

PLANT DESCRIPTION
Erected or spreading subshrub 50-80
cm high, with thick, fleshy leaves. En-
demic to Cyprus, restricted in fissures
of limestone cliffs along the Pentadak-
tylos mountain range (300-850 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Some rays become
dilated. Sclerenchyma cells in phloem
and in cortex. Sclereids in groups.
Phellem homogeneous, distinct in po-
larized light.

Xylem
As stem wood.

›T

Pith
Pith shape round. Medullary sheath
present. Heterogenous pith. Sclereids
present. Thick-walled parenchyma cells
Brassica hilarionis Post

present. Cells dimorphic. Cell content


present (dark staining substances). Pits
in transverse and in longitudinal cell
walls.

›T ›T

184
Stem xylem: 1 5 9 9.1 13 20 22 26 30 40.1 49 52.2 61 64 78 89 89.1 98 109 115
Twig bark: B1 B4 B6 B7 B19 B31 B33
Pith: P1 P2 P1 P3 P3.1 P3.3 P4 P5 P9 P9.1

Brassicaeae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands. Wood
diffuse-porous. Vessels solitary or in
radial multiples of 2 to 4 common.
Mean tangential diameter of early-
wood vessels less than 20 μm, 40-100
vessels per mm2. Parenchyma scanty
paratracheal, apotracheal parenchyma
in marginal or in seemingly marginal
bands, dark in polarized light. Rays
4-12 per mm. Groups of sieve tube into
the marginal parenchyma bands.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits scalariform and alternate, me-
dium (7-10 μm in diameter). Vessel-
ray pits with distinct borders, similar
to intervessel pits in size and shape
throughout the ray cell. Earlywood
vessel element length 50-100 μm. Fib-
ers with simple to minutely bordered
pits (libriform fibers). Helical thicken-
ings in ground tissue fibers. Rays with
procumbent, square and upright cells
mixed throughout the ray.

›T ›T

Tangential section
Rays commonly 4 to 10 seriate.
Brassica hilarionis Post

Wood basic density: 0,63 g/cm³

›T ›T

185
Erysimum kykkoticum Alziar et Hadjik.
Brassicaeae

PLANT DESCRIPTION
Subshrub up to 50 cm high, including
the inflorescence stalk. Leaves densely
covered with bifid hairs. A rare en-
demic species of Cyprus, which mostly
grows in crevices of igneous, diabase
rocks, occasionally on vertical banks
of abandoned tracks, of northern or
eastern aspects (250-450 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes in tangential rows. Cortex
consist of large and small thin-walled
parenchyma cells. Phellem homogene-
ous, distinct in polarized light.

Xylem
As stem wood.
Erysimum kykkoticum Alziar et Hadjik.

Pith
No slide available.

›T

186
Stem xylem: 1 5 9 13 20 24 39.1 40.1 50.2 52.2 58 61 69 89.2 117 133.2
Twig bark: B1 B4 B31 B33
Pith: No slide available.

Brassicaeae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands. Wood
diffuse-porous. Vessels predominantly
solitary. Vessels cell wall thick (more
than 2 μm). Mean tangential diameter
of earlywood vessels less than 20 μm.
More than 200 vessels per mm2. Gums
and other deposits in heartwood ves-
sels. Fibers thin- to thick-walled. Ring
shake. Included phloem: concentric
continuous.

›T

Radial section
Simple perforation plates. Inter-vessel
pits scalariform, minute (less than 4
μm in diameter). Earlywood vessel ele-
ment length 50-100 μm. Fibers with
simple to minutely bordered pits (libri-
form fibers).

Erysimum kykkoticum Alziar et Hadjik.


›T ›T

Tangential section
Wood rayless.

Wood basic density: 0,44 g/cm³

›T ›T

187
Buxus sempervirens L.
Common Box, Boxwood
Buxaceae

PLANT DESCRIPTION
Evergreen shrub up to 6 m high, with
glabrous, tetragonal shoots. Leaves
simple, 15-30x7-15 mm, dark green,
leathery. Exotic to Cyprus (150-1700
m alt.), indigenous to some Mediterra-
nean countries an central Europe.

TWIG ANATOMY
Bark
Groups of sieve tube present. Fibers
grouped. Cortex consist of large and
small thin-walled, unlignified, paren-
chyma cells. Epidermis distinct in
polarized light.

Xylem
As stem wood.

›T

Pith
Pith shape polygonal. Cells dimorphic.
Pits in transverse and in longitudinal
cell walls. Vascular bundles clearly sep-
arate. Tracheary elements of metax-
ylem in distinct radial rows.
Buxus sempervirens L.

›T ›T

188
Stem xylem: 1 5 9 14 15 21 24 30 40.2 50.1 53.2 60 62 70 76 78 86 97 100 107 108 115
Twig bark: B1 B9 B14 B14.3
Pith: P1.1 P4 P9 P9.1 P10.1 P12

Buxaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted latewood fibers. Wood
diffuse-porous. Vessels predominantly
solitary. Mean tangential diameter of
earlywood vessels 20-50 μm, 100-200
vessels per mm2. Fibers very thick-
walled. Axial parenchyma diffuse and
scanty paratracheal. Apotracheal pa-
renchyma in narrow bands or lines up
to three cells wide. Rays per mm 4-12.

›T ›T

Radial section
Scalariform perforation plates with
less than 10 bars. Inter-vessel pits
opposite, minute (less than 4 μm in
diameter). Vessels-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessels length more than
500 μm. Vascular and/or vasicentric
tracheids present. Fibers with distinctly
bordered pits (fiber tracheids). Body
ray cells procumbent with 2 to more
than 4 rows of square marginal cells.

›T ›T

Tangential section
Ray width predominantly 1 to 3 cells.
Rays with multiseriate portions as wide
as uniseriate portions.
Buxus sempervirens L.

Wood basic density: 0,82 g/cm³

›T ›T

189
Ceratonia siliqua L.
Carob Tree
Caesalpiniaceae

PLANT DESCRIPTION
Evergreen tree up to 10-15 m high,
with a broad crown. Leaves are 10 to
20 cm long, alternate and pinnate.
Indigenous to Cyprus, mostly a con-
stituent of maquis forests (0-600 m
alt.). It is widely cultivated for its edible
legumes. Widespread in the Mediterra-
nean from Portugal to Syria.

TWIG ANATOMY
Bark
Groups of sieve tube present. Distinct
rays dilatation. Sclerenchyma cells in
phloem and in cortex. Fibers in tan-
gential rows and scattered or irregu-
larly dispersed. Sclereids in tangential
rows or in groups. Prismatic crystals
present. Cell content in parenchyma
cells. Phellem homogeneous. Epider-
mis distinct in polarized light.

Xylem
First growth ring with few vessels and
very thick-walled fibers. Vessels in
radial multiples of 4 or more common.
›T
Rays predominantly uniseriate.

Pith
Pith shape polygonal. Cells dimorphic,
with small cells in lines. Heterogene-
ous pith. Unlignified cells present. Cell
content present (dark staining sub-
stances). Prismatic crystals and crystal
druses present. Pits in transverse and
Ceratonia siliqua L.

in longitudinal cell walls. Vascular bun-


dles clearly separate to not distinct.
Tracheary elements of metaxylem in
distinct radial rows.

›T ›T

190
Stem xylem: 1 5 9 9.1 11 13 22 25 30 41 45 49 53.1 56 58 61 65 69 70 77 79 83 96 97 105 106 116.1 136 141.1
Twig bark: B1 B6 B7 B12 B13 B17 B20 B28 B31 B33
Pith: P1.1 P3 P3.4 P4 P5 P6 P6.2 P9 P9.1 P10.1 P10.2 P12

Caesalpiniaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted latewood fibers. Wood
diffuse-porous. Vessels solitary, in
short radial multiples or in clusters.
Mean tangential diameter of early-
wood vessel lumina 50-100 μm, 40-
100 vessels per mm2. Vessels of two
distinct diameter classes. Tyloses with
thin walls common. Gums and other
deposits in heartwood vessels. Fibers
thick- to very thick-walled. Apotracheal
parenchyma diffuse in aggregates.
Paratracheal parenchyma vasicentric to
›T ›T
confluent. Rays 12-20 per mm.

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 200-
500 μm. Fibers with simple to minutely
bordered pits (libriform fibers). Septate
fibers present. All uniseriate rays with
square cells. Multiseriate rays com-
posed by procumbent cells with one
row of square marginal cells. Prismatic
crystals in axial parenchyma cells.

›T ›T

Tangential section
Ray width predominantly 1 to 3(4)
cells.
Ceratonia siliqua L.

Wood basic density: 0,61 g/cm³

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191
Celtis australis L.
Southern Nettle-Tree, Hackberry
Cannabaceae

PLANT DESCRIPTION
Deciduous tree up to 20 m high, with
silver-greyish bark. Flowers inconspicu-
ous green to yellowish; fruit a globose
fleshy dark brown drupe. Naturalized
on Cyprus, with a limited distribution,
although in some part it is locally com-
mon (0-1400 m alt.).

TWIG ANATOMY
Bark
Collapsed sieve tubes present. Some
rays become dilated. Sclerenchyma
cells in phloem and in cortex. Fibers
in tangential rows. Sclereids mostly
in groups. Prismatic crystals present.
Phellem homogeneous, distinct in po-
larized light.

Xylem
Vessels in radial multiples of 2 to 4.
Mean tangential diameter of earlywood
vessel lumina 50-100 μm. Ray width
predominantly 1 to 3 cells. Rays 4-12
per mm.
›T

Pith
Pith shape round. Pith partially disap-
pears in polarized light. Cells dimor-
phic. Prismatic crystals present. Pits of
two distinct size in transverse and in
longitudinal cell walls. Vascular bun-
dles clearly separate. Tracheary ele-
ments of metaxylem in distinct radial
Celtis australis L.

rows. Axial cells in regular rows (radial


section).

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192
Stem xylem: 1 3 6 9 11 13 22 26 31 36 42 49 52.3 56 61 69 70.2 79 83 85 96 99 102 103 107 114 136 136.1
Twig bark: B4 B6 B7 B9 B12 B15 B19 B20 B31 B33
Pith: P1.1 P1 P4 P6 P9 P9.1 P9.3 P10.1 P12 P13

Cannabaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
the difference in vessel size between
latewood and earlywood. Wood ring-
porous. Vessels solitary or in clusters.
Vessels arrangement in intra-annual
tangential bands. Mean tangential
diameter of earlywood vessel lumina
100-200 μm, 40-100 vessels per mm2.
Tyloses with thin walls common. Fib-
ers thin- to thick-walled. Tension wood
present. Axial parenchyma vasicentric
and confluent, parenchyma bands
more than three cells wide. Less than 4
rays per mm.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, medium size (7-10 μm
in diameter). Vessel-ray pits rounded
or angular with large apertures. Helical
thickenings in vessel elements present.
Earlywood vessel element length 100-
200 μm. Fibers with simple to minutely
bordered pits (libriform fibers). Body
ray cells procumbent with mostly 2-4
rows of square marginal cells. Prismat-
ic crystals present in ray parenchyma
cells.

›T ›T

Tangential section
Rays of two distance size: uniseriate
and more than 10 seriate. Larger rays
height more than 1 mm.
Celtis australis L.

Wood basic density: 0,65 g/cm³

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193
Celtis tournefortii Lam.
Oriental Nettle-Tree
Cannabaceae

PLANT DESCRIPTION
Deciduous tree or shrub, 2-8 m high.
Very similar to Celtis australis, differ-
ing in its yellow brown fruits and gla-
brous leaves. Indigenous in the eastern
Mediterranean countries, it is very rare
on Cyprus (600-1400 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclerenchyma cells
in phloem and in cortex. Sclereids in
tangential rows and in groups. Prismat-
ic crystals present. Phellem homogene-
ous, distinct in polarized light.

Xylem
Rays predominantly 1 to 3 cells wide.
Rays 4-12 per mm.

›T

Pith
Pith shape round. Cells dimorphic.
Prismatic crystals present. Pits in trans-
verse and in longitudinal cell walls.
Celtis tournefortii Lam.

Vascular bundles clearly separate.


Tracheary elements of metaxylem in
distinct radial rows. Axial cells in regu-
lar rows (radial section).

›T ›T

194
Stem xylem: 1 3 6 11 13 22 25 30 36 41 49 52.3 61 69 70.2 77 79 83 96 98 106 115 136 136.1
Twig bark: B1 B4 B7 B17 B19 B20 B31 B33
Pith: P1 P4 P6 P9 P10.1 P12 P13

Cannabaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by the
difference in vessel size between late-
wood and earlywood. Wood ring-po-
rous. Vessels arrangement in intra-an-
nual tangential bands. Vessel clusters
common. Mean tangential diameter of
earlywood vessel lumina 50-100 μm,
40-100 vessels per mm2. Fibers thin- to
thick-walled. Tension wood present.
Axial parenchyma vasicentric to conflu-
ent. Apotracheal parenchyma diffuse in
aggregates. Rays 4-12 per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Helical thickenings in vessel elements
present. Earlywood vessel element
length 100-200 μm. Fibers with simple
to minutely bordered pits (libriform
fibers). Body ray cells procumbent with
one row of upright and square margin-
al cells. Prismatic crystals in ray paren-
chyma cells.

›T ›T

Tangential section
Rays uniseriate and up to 10 seriate.
Celtis tournefortii Lam.

Wood basic density: 0,69 g/cm³

›T ›T

195
Capparis spinosa L. (sensu Meikle)
Caper
Capparaceae

PLANT DESCRIPTION
Much-branched, deciduous, thorny
shrub up to 1,5 m high. Leaves sim-
ple, alternate, ovate. It occurs on
rocky slopes, in field margins, maquis,
garigue and wasteland (0-900 m alt.).
Also indigenous to other Mediterra-
nean countries.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Only some rays
become dilated. Fibers and sclereids
present, scattered or irregularly dis-
persed. Prismatic crystals present.
Phellem distinct in polarized light.

Xylem
Vessels predominantly solitary. Rays
predominantly uniseriate. Less than 20
rays per mm. Few prismatic crystals.
Capparis spinosa L. (sensu Meikle)

›T

Pith
Pith shape round. Pith not visible in
polarized light. Medullary sheath pre-
sent. Cells dimorphic. Prismatic crys-
tals present. Pits of two distinct size
in transverse and in longitudinal cell
walls. Vascular bundles clearly sepa-
rate. Tracheary elements of metaxylem
in distinct radial rows. Axial cells in
regular rows (radial section).

›T ›T

196
Stem xylem: 1 3 4 9 9.1 10 13 22 24 30 40.2 45 49 52.3 60 61 69 78 96.1 97 106 109 116.2
Twig bark: B1 B4 B6 B9 B13 B15 B18 B20 B14.33
Pith: P1.1 P1 P2 P4.1 P6 P9 P9.1 P9.3 P10.1 P12 P13

Capparaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by the
difference in vessel size between late-
wood and earlywood and by marginal
parenchyma cells. Wood ring-porous
to semi-ring-porous. Vessels predomi-
nantly solitary or in radial multiples
of 2 to 6 elements. Mean tangential
diameter of earlywood vessels 20-50
μm, 40-100 vessels per mm2. Vessels
of two distinct diameter classes. Fibers
thin- to thick-walled. Axial parenchyma
scanty paratracheal. More than 20 rays
per mm.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, minute (less than 4
μm in diameter). Vessel-ray pits with
distinct borders, similar to interves-
sel pits in size and shape throughout
the ray cell. Earlywood vessel element
length 100-200 μm. Vascular and/or
vasicentric tracheids present. Fibers
with simple to minutely bordered pits
(libriform fibers). Body ray cells mostly
procumbent with one row of upright
and square marginal cells. Uniseriate
rays with procumbent, square and up-
right cells mixed throughout the ray.

Capparis spinosa L. (sensu Meikle)


›T ›T

Tangential section
Rays predominantly 1 to 4 cells.

Wood basic density: 0,45 g/cm³

›T ›T

197
Lonicera etrusca Santi
Estruscan Honeysuckle
Caprifoliaceae

PLANT DESCRIPTION
Deciduous climber with stems up to
4 m long. Young shoots usually pur-
ple, mainly glabrous. Leaves opposite,
simple, variously petiolate. Indigenous
to Cyprus, growing in thickets, pine
forests and rocky places (300-1500 m
alt.). It is also native in other Mediter-
ranean countries.

TWIG ANATOMY
Bark
Sclerenchyma cells in phloem and in
cortex. Fibers in tangential rows.
Phellem homogeneous, distinct in po-
larized light. Lignified cells in phellem.

Xylem
Mean tangential diameter of earlywood
vessels 20-50 μm. Rays predominantly
uniseriate with upright and squared
cells only. More than 20 rays per mm.

›T

Pith
Pith not visible in polarized light. Pith
shape round. Medullary sheath pre-
sent. Heterogeneous pith. Unlignified
cells present. Cells dimorphic. Pits
Lonicera etrusca Santi

in transverse and in longitudinal cell


walls. Pits of two distinct size. Vas-
cular bundles clearly separate to not
distinct. Tracheary elements of metax-
ylem in distinct radial rows.

›T ›T

198
Stem xylem: 1 5 9 9.1 13 22 25 30 36 40.2 50.2 53.1 58 60 62 69 70.3 75 76 96 97 105 109 116.1
Twig bark: B7 B12 B31 B33 B34
Pith: P1 P2 P3 P3.4 P4 P9 P9.1 P9.3 P10.1 P10.2 P12

Caprifoliaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood diffuse-porous. Vessels
solitary and in radial multiples of 2 to
4 common. Mean tangential diameter
of earlywood vessels 20-50 μm, more
than 200 vessels per mm2. Gums and
other deposits in heartwood vessels.
Fibers thin- to thick-walled. Radial flat
marginal fibers. Axial parenchyma ab-
sent or extremely rare or not to recog-
nizable, few scatter apotracheal diffuse
parenchyma cells present. Rays 12-20
per mm.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Helical thickenings in vessel elements
present. Earlywood vessel element
length 200-500 μm. Vascular and/or
vasicentric tracheids present. Fibers
with distinctly bordered pits (fiber trac-
heids). All ray cells upright and square.
Rays with procumbent, square and
upright cells mixed throughout the ray.

›T ›T

Tangential section
Rays uniseriate and up to 3 cells wide.
Lonicera etrusca Santi

Wood basic density: 0,49 g/cm³

›T ›T

199
Lonicera japonica Thunb.
Japanese Honeysuckle
Caprifoliaceae

PLANT DESCRIPTION
Climbing species with stems up to 6 m
long. Young shoot hairy. Leaves oppo-
site, oblong to ovate. Exotic to Cyprus,
occurring in gardens and hedges (0-
500 m alt.). Cultivated in Mediterrane-
an countries. Indigenous to south-east
Asia.

Photo: Hadjikyriakou
TWIG ANATOMY
Bark
Groups of sieve tube present.
Sclereids scattered or irregularly dis-
persed. Crystal druses present. Cell
content in parenchyma cells. Phellem
homogeneous, consist of thin walled
cells. Phellem distinct in polarized
light.

Xylem
Mean tangential diameter of earlywood
vessels 20-50 μm. Rays predominantly
uniseriate with upright and squared
cells only. More than 20 rays per mm.
›T

Pith
Pith shape square to roundish.
Medullary sheath present. Heterogene-
Lonicera japonica Thunb.

ous pith. Unlignified cells in the centre


of the pith, lignified parenchyma cells
at the periphery. Pits in transverse
and in longitudinal cell walls. Vascular
bundles clearly separate. Axial cells in
regular rows (radial section).

›T ›T

200
Stem xylem: 1 3 4 11 13 22 25 31 36 41 45 49 53.1 60 62 69 70.3 78 96 98 108 116.1 136 136.1
Twig bark: B1 B18 B22 B28 B31 B33
Pith: P1.4 P2 P3.4 P9 P10.1 P13

Caprifoliaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood semi-ring to ring-porous.
Vessel clusters common. Vessels of
two distinct diameter classes. Mean
tangential diameter of earlywood ves-
sel lumina 50-100 μm, 40-100 vessels
per mm2. Fibers thin- to thick-walled.
Radial flat marginal fibers. Axial paren-
chyma scanty paratracheal. Rays 12-20
per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits rounded or
angular with large apertures. Helical
thickenings in vessel elements present.
Earlywood vessel element length 200-
500 μm. Vascular and/or vasicentric
tracheids present. Fibers with distinctly
bordered pits (fiber tracheids). Body
ray cells procumbent with over 4 rows
of upright and square marginal cells.
Prismatic crystals in ray parenchyma
cells.

›T ›T

Tangential section
Rays uniseriate to 10 seriate.
Lonicera japonica Thunb.

Wood basic density: 0,56 g/cm³

›T ›T

201
Dianthus cyprius A.K. Jacks. et Turrill
Cyprus Pink
Caryophyllaceae

PLANT DESCRIPTION
Suffrutticose perennial, with sparse,
pendulous and often curved branches,
up to 1 m long. Endemic to Cyprus
growing on limestone rocks and cliffs
on the Pentadaktylos range (350-750
m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube present. Fibers in
tangential rows and grouped. Crystal
sand and druses present. Layered phel-
loderm.

Xylem
As stem wood.
Dianthus cyprius A.K. Jacks. et Turrill

›T

Pith
Pith not visible in polarized light. Pith
shape round. Medullary sheath pre-
sent. Heterogeneous pith. Thick-walled
and unlignified cells present. Cells
dimorphic. Crystal sand present. Vas-
cular bundles not distinct.

›T

202
Stem xylem: 1 5 9.1 13 22 24 39.1 40.1 50.1 52.2 61 69 78 89 89.1 117
Twig bark: B1 B12 B14 B22 B23 B29
Pith: P1 P2 P3 P3.3 P3.4 P4 P6.4 P10.2

Caryophyllaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
a marginal parenchyma band. Wood
diffuse-porous. Vessels in radial mul-
tiples of 2 to 4 common. Vessels cell
wall thick (more than 2 μm). Mean tan-
gential diameter of earlywood vessels
less than 20 μm, 100-200 vessels per
mm2. Fibers thin- to thick-walled. Pa-
renchyma scanty paratracheal. Apotra-
cheal parenchyma in marginal or in
seemingly marginal bands. Parenchyma
marginal thin-walled, dark in polarized
light.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, minute (less than 4 μm
in diameter). Earlywood vessel element
length 50-100 μm. Fibers with simple
to minutely bordered pits (libriform
fibers).

Tangential section Dianthus cyprius A.K. Jacks. et Turrill


Wood rayless.

Wood basic density: 0,43 g/cm³

›T

203
Silene fruticosa L.
Caryophyllaceae

PLANT DESCRIPTION
A loosely tufted subshrub, 15-50 cm
high. A native of Cyprus growing in
rock crevices and on old walls, with a
limited distribution, although locally
common in some areas (0-600 m alt.).
It is also found in Malta, Sicily, Greece
and Lybia.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Crystal druses
present. Phellem homogeneous, dis-
tinct in polarized light. Lignified cells
in phellem.

Xylem
Mean tangential diameter of earlywood
vessels less than 20 μm. Earlywood
vessel element length 100-200 μm.

›T

Pith
Pith not visible in polarized light.
Pith shape round. Heterogenous pith.
Thick-walled and unlignified paren-
chyma cells present. Cells dimorphic.
Crystal druses present. Pits in trans-
verse and in longitudinal cell walls.
Vascular bundles not distinct.
Silene fruticosa L.

›T

204
Stem xylem: 1 5 9 13 20 22 26 36 39.1 40.2 50.1 52.2 61 69 78 89 89.1 117
Twig bark: B1 B4 B22 B31 B33 B34
Pith: P1 P3 P3.3 P3.4 P4 P6.2 P9 P9.1 P10.2

Caryophyllaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
a marginal parenchyma band. Wood
diffuse-porous. Vessels predominantly
solitary. Vessels cell wall thick (more
than 2 μm). Mean tangential diameter
of earlywood vessels 20-50 μm, 100-
200 vessels per mm2. Fibers thin- to
thick-walled. Axial parenchyma scanty
paratracheal. Apotracheal parenchyma
in marginal or in seemingly marginal
bands, with thin-walled cells, dark in
polarized light.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits scalariform and alternate, medium
(7-10 μm in diameter). Helical thicken-
ings in vessel elements. Earlywood ves-
sel element length 50-100 μm. Fibers
with simple to minutely bordered pits
(libriform fibers).

›T ›T

Tangential section
Wood rayless.
Silene fruticosa L.

Wood basic density: 0,38 g/cm³

›T ›T

205
Silene galatea Boiss.
Caryophyllaceae

PLANT DESCRIPTION
Subshrub 20-50 m high, with glabrous
suberect stems. Endemic to Cyprus,
thriving on dry rocky slopes in Pine
forests, garigue and screes (750-1200
m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube in tangential
rows. Collapsed sieve tubes present.
Prismatic crystals present. Phellem
consist of thin-walled cells.

Xylem
As stem wood.

›T

Pith
Pith not visible in polarized light.
Unlignified cells present. Pith shape
round. Cells dimorphic. Vascular bun-
dles not distinct.
Silene galatea Boiss.

›T ›T

206
Stem xylem: 2 5 9 13 20 25 39.1 40.1 50.2 53.1 61 69 79.1 117
Twig bark: B1 B2 B4 B20
Pith: P0.1 P1 P3.1 P4 P10.2

Caryophyllaceae
STEM ANATOMY
Transverse section
Growth ring indistinct or absent. Wood
diffuse-porous. Vessels predominantly
solitary. Vessels cell wall thick (more
than 2 μm). Mean tangential diameter
of earlywood vessels less than 20 μm.
More than 200 vessels per mm2. Fib-
ers thin- to thick-walled, in tangentially
elongated groups. Parenchyma perva-
sive.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits scalariform, small (4-7 μm in diam-
eter). Earlywood vessel element length
200-500 μm. Fibers with simple to mi-
nutely bordered pits (libriform fibers).

Tangential section
Wood rayless.
Silene galatea Boiss.

Wood basic density: 0,63 g/cm³

›T

207
Arthrocnemum macrostachyum (Moric.) Delponte et Moris
Chenopodiaceae

PLANT DESCRIPTION
Erect, much-branched subshrub 20-70
cm high. Young stems fleshy, older
stems woody. Leaves fleshy, scale-like.
An indigenous halophyte, occurring on
edges of salt lakes and salt marshes
(0-150 m alt.). Indigenous of the Medi-
terranean region, Red Sea, Atlantic
islands and south to Senegal.

TWIG ANATOMY
Arthrocnemum macrostachyum (Moric.) Delponte et Moris

Bark
Crystal druses present. The cortex
consist of large and small parenchyma
cells. The phellem consist of thick-
walled cork cells, distinct in polarized
light.

Xylem
As stem wood.

›T

Pith
Pith not visible in polarized light.
Pith shape round. Cells dimorphic. Vas-
cular bundles not distinct.

›T ›T

208
Stem xylem: 1 5 9 11 13 22 24 39.1 40.1 50.1 52.2 61 69 70 78 117 133.1
Twig bark: B22 B31 B33
Pith: P0.1 P1 P4 P10.2

Chenopodiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood diffuse-porous. Vessels
solitary and in clusters. Vessels cell
wall thick (more than 2 μm). Mean tan-
gential diameter of earlywood vessels
less than 20 μm, 100-200 vessels per
mm2. Fibers thick- to very thick-walled.
Axial parenchyma scanty paratracheal.
Included phloem: concentric arranged
single vascular bundles.

›T ›T

Radial section

Arthrocnemum macrostachyum (Moric.) Delponte et Moris


Simple perforation plates. Inter-vessel
pits alternate, minute (less than 4 μm
in diameter). Earlywood vessel element
length 50-100 μm. Fibers with simple
to minutely bordered pits (libriform
fibers).

›T ›T

Tangential section
Wood rayless.

Wood basic density: 0,67 g/cm³

›T ›T

209
Atriplex halimus L.
Shrubby Saltbush, Tree Purslane
Chenopodiaceae

PLANT DESCRIPTION
Erect shrub up to 2 m high, with sil-
very-grey leaves. Bark peeling off in
narrow longitudinal strips. Indigenous
species occurring on sandy and rocky
seashores and sometimes on dry sites
inland (0-350 m alt.). Widespread in
the Mediterranean region.

TWIG ANATOMY
Bark
Groups of sieve tube present. Fibers in
tangential rows. Prismatic crystals pre-
sent. Anatomy difficult to interpret.

Xylem
As stem wood.

›T

Pith
Pith shape round. Cells dimorphic.
Prismatic crystals present. Crystal
druses present. Pits in transverse and
in longitudinal cell walls. Vascular
bundles in the pith. Vascular bundles
clearly separate. Axial cells in regular
Atriplex halimus L.

rows (radial section).

›T ›T

210
Stem xylem: 1 5 11 13 22 25 39.1 40.2 45 50.1 52.2 61 69 70 78 117 120 133.1 136
Twig bark: B1 B12 B20
Pith: P1 P4 P6 P6.2 P9 P9.1 P10 P10.1 P13

Chenopodiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted and thick-walled late-
wood fibers. Wood diffuse-porous.
Vessel clusters common. Vessels cell
wall thick (more than 2 μm). Mean tan-
gential diameter of earlywood vessels
20-50 μm, 100-200 vessels per mm2.
Vessels of two distinct diameter class-
es. Fibers thick- to very thick-walled.
Axial parenchyma scanty paratracheal.
Included phloem: concentric arranged
single vascular bundles.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in diam-
eter). Earlywood vessel element length
50-100 μm. Fibers with simple to mi-
nutely bordered pits (libriform fibers).
Prismatic crystals present.

›T ›T

Tangential section
Wood rayless. Axial parenchyma and
vessels storied.
Atriplex halimus L.

Wood basic density: 0,65 g/cm³

›T ›T

211
Atriplex portulacoides L.
= Halimione portulacoides (L.) Aellen
Chenopodiaceae

PLANT DESCRIPTION
Sprawling much-branched subshrub
1,5 m high. Shoots angular in cross
section. Leaves opposite, oblong-ellip-
tic, greyish-green. Indigenous species
occurring on edges of salt lakes and
sea shores (0 m alt.). Indigenous to
western Europe and the Mediterranean
region.

Photo: Hadjikyriakou
TWIG ANATOMY
Bark
Fibers scattered or irregularly dis-
persed. Crystal druses present. The
cortex consist of large and small
parenchyma cells. Phellem homogene-
ous, distinct in polarized light. Ligni-
fied cells in phellem.

Xylem
As stem wood.

›T

Pith
Pith not visible in polarized light.
Pith shape round. Heterogenous pith.
Thick-walled parenchyma cells and
Atriplex portulacoides L.

unlignified cells present. Pits in trans-


verse and in longitudinal cell walls.
Vascular bundles in the pith. Vascular
bundles clearly separate. Tracheary el-
ements of metaxylem in distinct radial
rows.

›T ›T

212
Stem xylem: 1 5 9 11 13 22 24 39.1 40.1 45 50.2 52.3 61 69 70 78 117 133.1
Twig bark: B13 B22 B31 B33 B34
Pith: P0.1 P1 P3 P3.3 P3.4 P9 P9.1 P10 P10.1 P12

Chenopodiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood diffuse-porous. Vessels
solitary and in clusters common. Ves-
sels cell wall thick (more than 2 μm).
Mean tangential diameter of earlywood
vessels less than 20 μm. More than
200 vessels per mm2. Vessels of two
distinct diameter classes. Fibers thick-
to very thick-walled. Axial parenchyma
scanty paratracheal. Included phloem:
concentric arranged single vascular
bundles.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, minute (less than 4 μm
in diameter). Earlywood vessel element
length 100-200 μm. Fibers with simple
to minutely bordered pits (libriform
fibers).

›T ›T

Tangential section
Wood rayless.
Atriplex portulacoides L.

Wood basic density: 0,55 g/cm³

›T ›T

213
Atriplex semibaccata R.Br.
Creeping Saltbush, Australian Saltbush
Chenopodiaceae

PLANT DESCRIPTION
Prostrate or sprawling subshrub with
stems up to 50 cm long. Leaves usually
opposite, ovate or narrowly oblong. A
naturalised species growing on waste
ground and garigue and on brackish
ground inland (30-250 m alt.). A native
of Australia, probably introduced to
Cyprus as a fodder plant.

TWIG ANATOMY
Bark
Phloem uniform. Prismatic crystals
present. The cortex consist of large
and small parenchyma cells. Phellem
consist of large and small cork cells.
Epidermis distinct in polarized light.

Xylem
As stem wood.

›T

Pith
Pith not visible in polarized light.
Heterogenous pith. Thick-walled paren-
Atriplex semibaccata R.Br.

chyma cells present. Cells dimorphic.


Crystal druses present. Pits in trans-
verse and in longitudinal cell walls.
Pits grouped and of two distinct size.
Vascular bundles in the pith. Vascular
bundles clearly separate.

›T ›T

214
Stem xylem: 1 5 11 13 22 24 39.1 40.2 45 50.1 52.2 58 61 69 70 78 117 133.1 133.2 136
Twig bark: B20 B8 B33
Pith: P3 P3.3 P4 P6.2 P9 P9.1 P9.2 P9.3 P10 P10.1

Chenopodiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood diffuse-porous. Vessels
in clusters common. Vessels cell wall
thick (more than 2 μm). Vessels of
two distinct diameter classes, mean
tangential diameter of vessels 20-50
μm, 100-200 vessels per mm2. Gums
and other deposits in heartwood ves-
sels. Fibers thick- to very thick-walled.
Axial parenchyma scanty paratracheal.
Included phloem: concentric arranged
single vascular bundles, sometimes
connected in transverse section.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, minute (less than 4 μm
in diameter). Earlywood vessel element
length 50-100 μm. Fibers with simple
to minutely bordered pits (libriform fib-
ers). Prismatic crystals present.

›T ›T

Tangential section
Wood rayless.
Atriplex semibaccata R.Br.

Wood basic density: 0,40 g/cm³

›T ›T

215
Halocnemum strobilaceum (Pall.) M.Bieb.
Chenopodiaceae

PLANT DESCRIPTION
Erect much-branched subshrub up to
80 cm high. Young shoots fleshy, glau-
cous, with cylindrical jointed branches.
An indigenous halophyte on Cyprus,
occurring on edges of salt lakes and
salt marshes (0 m alt.). Also indig-
enous to the Mediterranean region and
eastward to Asia.

TWIG ANATOMY
Bark
Phloem uniform. Anatomy difficult to
interpret.

Xylem
As stem wood.
Halocnemum strobilaceum (Pall.) M.Bieb.

›T

Pith
Pith not visible in polarized light. Pith
shape polygonal. Heterogeneous pith.
Unlignified cells present. Cell dimor-
phic. Pits in transverse and in longitu-
dinal cell walls. Vascular bundles in the
pith. Vascular bundles clearly separate.
Tracheary elements of metaxylem in
distinct radial rows.

›T ›T

216
Stem xylem: 2 5 9.1 11 13 22 25 36 39.1 40.1 49 52.2 61 78 117 120 133.1
Twig bark: B8
Pith: P0.1 P1.1 P3.4 P4 P9 P9.1 P10 P10.1 P12

Chenopodiaceae
STEM ANATOMY
Transverse section
Growth rings indistinct or absent.
Wood diffuse-porous. Vessels in radial
multiples of 2 to 4 common. Vessel
clusters common. Vessels cell wall
thick (more than 2 μm). Mean tangen-
tial diameter of vessels less than 20
μm, 40-100 vessels per mm2. Axial pa-
renchyma scanty paratracheal. Includ-
ed phloem: concentric arranged single
vascular bundles.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in diam-
eter). Helical thickenings in vessel ele-
ments present. Vessel element length
50-100 μm. Fibers with simple to mi-
nutely bordered pits (libriform fibers).

Halocnemum strobilaceum (Pall.) M.Bieb.


›T ›T

Tangential section
Wood rayless. Axial parenchyma sto-
ried.

Wood basic density: 1,00 g/cm³

›T ›T

217
Noaea mucronata (Forssk.) Schweinf. et Asch.
Thorny Saltwort
Chenopodiaceae

PLANT DESCRIPTION
Erect or sprawling, much-branched
subshrub up to 60 cm high. Indig-
enous to Cyprus, occurring on dry pas-
tures, costal garigue, sand dunes and
rocky slopes (0-650 m alt.). Indigenous
to the eastern Mediterranean region,
north Africa and eastward to central
Asia.

TWIG ANATOMY
Bark
Phloem uniform. Layered phelloderm.
Lignified cells in phellem. Cortex con-
sist of large thin-walled cells. Anatomy
difficult to interpret.

Xylem
As stem wood.
Noaea mucronata (Forssk.) Schweinf. et Asch.

›T

Pith
Pith shape round. Heterogeneous pith.
Thick-walled parenchyma cells present.
Cells dimorphic. Pits in transverse and
in longitudinal cell walls. Vascular bun-
dles clearly separate.

›T ›T

218
Stem xylem: 1 5 9 11 13 24 39.1 40.1 50.1 52.3 61 69 70 78 117 133.1
Twig bark: B8 B34 B35
Pith: P1 P3.3 P4 P9 P9.1 P10.1

Chenopodiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood diffuse-porous. Vessels
solitary and in clusters common. Ves-
sels cell wall thick (more than 2 μm).
Mean tangential diameter of earlywood
vessels less than 20 μm, 100-200
vessels per mm2. Fibers thick- to very
thick-walled. Axial parenchyma scanty
paratracheal. Included phloem: concen-
tric arranged single vascular bundles.

›T ›T

Radial section
Simple perforation plates. Intervessel
pits minute (less than 4 μm in diam-
eter). Earlywood vessel element length
100-200 μm. Fibers with simple to mi-
nutely bordered pits (libriform fibers).

Noaea mucronata (Forssk.) Schweinf. et Asch.


›T ›T

Tangential section
Wood rayless.

Wood basic density: 0,86 g/cm³

›T ›T

219
Salicornia fruticosa (L.) L.
Glasswort
Chenopodiaceae

PLANT DESCRIPTION
Erect, much branched subshrub 20-70
cm high. Young stems fleshy, older
stems woody, with jointed branches.
Leaves opposite, fleshy, scale-like. An
indigenous halophyte, occurring on
edges of salt lakes and salt marshes
(0-10 m alt.). Indigenous to the Medi-
terranean and the Atlantic islands.

TWIG ANATOMY
Bark
Groups of sieve tube present. Phloem
uniform. Cell content in phloem and
cortex cells. Phellem homogeneous,
consist of thick-walled cells, distinct in
polarized light.

Xylem
As stem wood.

›T

Pith
Pith not visible in polarized light. Pith
shape round. Heterogeneous pith.
Salicornia fruticosa (L.) L.

Unlignified cells present. Vascular bun-


dles clearly separate.

›T ›T

220
Stem xylem: 2 5 9 11 13 22 24 39.1 40.1 49 53.1 61 70 78 117 120 133.1
Twig bark: B1 B8 B28 B31 B33
Pith: P0.1 P1 P3.4 P10.1

Chenopodiaceae
STEM ANATOMY
Transverse section
Growth ring indistinct or absent. Wood
diffuse-porous. Vessels solitary and in
clusters. Vessels cell wall thick (more
than 2 μm). Mean tangential diameter
of vessels less than 20 μm, 40-100 ves-
sels per mm2. Fibers very thick-walled.
Axial parenchyma scanty paratracheal.
Included phloem: concentric arranged
single vascular bundles.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, minute (less than 4 μm
in diameter). Vessels length 200-500
μm. Fibers with simple to minutely bor-
dered pits (libriform fibers).

›T ›T

Tangential section
Wood rayless. Axial parenchyma and
vessel elements storied.
Salicornia fruticosa (L.) L.

Wood basic density: 0,87 g/cm³

›T ›T

221
Sarcocornia perennis (Mill.) A.J. Scott
Perennial Glasswort
Chenopodiaceae

PLANT DESCRIPTION
Subshrub 10-50 m high with creeping,
fleshy stems rooting at nodes. Leaves
opposite, fleshy. And indigenous halo-
phyte of Cyprus, occurring on edges
of salt lakes and salt marshes (0-100
m). Indigenous to the Mediterranean
region, southern and western Europe,
tropical and southern Africa.

TWIG ANATOMY
Bark
Phloem uniform. Prismatic crystals pre-
sent. Phellem homogeneous. Epidermis
distinct in polarized light. Anatomy
difficult to interpret.

Xylem
As stem wood.
Sarcocornia perennis (Mill.) A.J. Scott

›T

Pith
Pith not visible in polarized light. Pith
star shape. Heterogeneous pith. Unlig-
nified cells present. Pits in transverse
and in longitudinal cell walls. Vascular
bundles not distinct.

›T ›T

222
Stem xylem: 2 5 9 11 13 22 24 39.1 40.1 50.1 52.2 61 69 70 78 117 120 133.1 133.2
Twig bark: B20 B8 B31 B33
Pith: P1 P3.4 P9 P10.2

Chenopodiaceae
STEM ANATOMY
Transverse section
Growth ring indistinct or absent. Wood
diffuse-porous. Vessels solitary or in
clusters. Vessels cell wall thick (more
than 2 μm). Mean tangential diameter
of earlywood vessels less than 20 μm,
100-200 vessels per mm2. Fibers thick-
to very thick-walled. Axial parenchyma
scanty paratracheal. Included phloem:
concentric arranged single vascular
bundles, sometimes concentric con-
tinuous.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, minute (less than 4 μm
in diameter). Vessel element length 50-
100 μm. Fibers with simple to minutely
bordered pits (libriform fibers).

Sarcocornia perennis (Mill.) A.J. Scott


›T ›T

Tangential section
Wood rayless. Axial parenchyma and
vessel elements storied.

Wood basic density: 0,65 g/cm³

›T ›T

223
Suaeda aegyptiaca (Hasselq.) Zohary
Aegyptian Sea-Blite
Chenopodiaceae

PLANT DESCRIPTION
Erect, much branched subshrub up to
50 cm high, with fleshy leaves. A rare
indigenous halophyte of Cyprus, oc-
curring on edges of salt lakes and salt
marshes (0 m alt.). It is indigenous
also to Libya, Egypt, Israel and east-
ward to Iran, Afghanistan, Pakistan and
southwards to Sudan.

TWIG ANATOMY
Bark
Phloem uniform. Crystal druses and
crystal sand present. The cortex con-
sist of large and small parenchyma
cells. Lignified cells in phellem.

Xylem
As stem wood.
Suaeda aegyptiaca (Hasselq.) Zohary

›T

Pith
Pith shape round. Cells dimorphic. Pits
in transverse and in longitudinal cell
walls. Vascular bundles in the pith.
Vascular bundles clearly separate.

›T ›T

224
Stem xylem: 2 5 9.1 10 13 22 25 30 39.1 40.2 50.1 53.1 61 69 78 79 97 109 115 120 133.1
Twig bark: B22 B23 B8 B34
Pith: P1 P4 P9 P9.1 P10 P10.1

Chenopodiaceae
STEM ANATOMY
Transverse section
Growth ring indistinct or absent. Wood
diffuse-porous. Vessels in radial mul-
tiples of 2 to 4 or more. Vessels cell
wall thick (more than 2 μm). Mean
tangential diameter of earlywood ves-
sels 20-50 μm, 100-200 vessels per
mm2. Fibers thin- to thick-walled. Axial
parenchyma scanty paratracheal to va-
sicentric. Rays 4-12 per mm. Included
phloem: concentric arranged single
vascular bundles.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 200-
500 μm. Fibers with simple to minutely
bordered pits (libriform fibers). Rays
with procumbent, square and upright
cells mixed throughout the ray.

Suaeda aegyptiaca (Hasselq.) Zohary


›T ›T

Tangential section
Ray width predominantly 1 to 3 cells.
Axial parenchyma storied.

Wood basic density: 0,49 g/cm³

›T ›T

225
Suaeda vera Forssk.
True Sea-Blite
Chenopodiaceae

PLANT DESCRIPTION
Evergreen, erect, much-branched shrub
up to 80 cm high, with fleshy leaves.
An indigenous halophyte, occurring on
sandy coasts, edges of salt lakes and
rocky slopes (0-30 m alt.). Indigenous
to the southern and western Europe,
the Mediterranean region and the At-
lantic islands.

TWIG ANATOMY
Bark
The phloem consist of large and small
parenchyma cells. Phellem with large
and small thin-walled cork cells.

Xylem
As stem wood.

›T

Pith
Pith shape round. Cells dimorphic.
Pits in longitudinal and transverse cell
walls. Vascular bundles in the pith,
clearly separated.
Suaeda vera Forssk.

›T ›T

226
Stem xylem: 1 5 9 11 13 22 25 36 39.1 40.1 50.1 52.2 58 61 69 78 83 85 89 117 133.1
Twig bark: B8
Pith: P1 P4 P9 P9.1 P10 P10.1

Chenopodiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted and thick-walled late-
wood fibers. Wood diffuse-porous.
Vessels solitary and in clusters. Vessels
cell wall thick (more than 2 μm). Mean
tangential diameter of earlywood ves-
sels less than 20 μm, 100-200 vessels
per mm2. Gums and other deposits
in heartwood vessels. Fibers thin- to
thick-walled. Axial parenchyma some-
times scanty paratracheal, often con-
fluent and in bands more than three
cells wide. Apotracheal parenchyma
in marginal or in seemingly marginal
›T ›T
bands. Included phloem: concentric ar-
ranged single vascular bundles.
Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Helical thickenings in vessel
elements present. Earlywood vessel
element length 50-100 μm. Fibers with
simple to minutely bordered pits (libri-
form fibers).

›T ›T

Tangential section
Wood rayless.
Suaeda vera Forssk.

Wood basic density: 0,62 g/cm³

›T ›T

227
Cistus creticus L. subsp. creticus
Rockrose, Cretan Rockrose
Cistaceae

PLANT DESCRIPTION
Shrub up to 150 cm high. Young shoots
and leaves covered with glandular and
glandular hairs. It is indigenous to cen-
tral and eastern Mediterranean coun-
tries as far as Palestine. Very common
on Cyprus, growing on rocky slopes,
forests and shrublands (0-1800 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube present. Scle-
renchyma cells in phloem and in cor-
tex. Fibers in tangential rows. Crystal
druses present.

Xylem
Wood semi-ring-porous to diffuse-
porous.

›T
Cistus creticus L. subsp. creticus

Pith
Pith shape round. Heterogeneous pith.
Thick-walled parenchyma cells. Cells
dimorphic. Crystal druses present. Pits
in longitudinal cell walls (radial sec-
tion). Vascular bundles clearly separate
to not distinct.

›T ›T

228
Stem xylem: 1 4 9 13 20.1 22 25 30 40.2 45 49 58 62 70 76 96 97 103 109 116.2
Twig bark: B1 B7 B9 B12 B22
Pith: P1 P3.3 P4 P6.2 P9 P10.1 P10.2

Cistaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted latewood fibers and by
marginal parenchyma bands. Wood
semi-ring-porous. Vessels predomi-
nantly solitary. Mean tangential diam-
eter of earlywood vessels 20-50 μm,
40-100 vessels per mm2. Vessels of
two distinct diameter classes. Gums
and other deposits in heartwood ves-
sels. Fibers very thick-walled. Axial
parenchyma diffuse. More than 20 rays
per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits pseudoscalariform to reticulate
and alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Fibers with distinctly bordered pits
(fiber tracheids). Rays with procum-
bent, square and upright cells mixed
throughout the ray.

›T ›T

Tangential section Cistus creticus L. subsp. creticus


Rays of two distinct sizes: uniseriate
and up to 3 cells wide.

Wood basic density: 0,71 g/cm³

›T ›T

229
Cistus ladanifer L.
Gum Cistus, Laudanum
Cistaceae

PLANT DESCRIPTION
Shrub up to 2 m high. Young shoots
viscid, with aromatic resin. Leaves
opposite, narrowly lanceolate, upper
surface leathery, glabrous, lower sur-
face grey-tomentose. Native to western
Mediterranean basin, it is a rare natu-
ralized species in Cyprus. It grows in
open pine forests (250-800 m alt.)

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Some rays become
dilated. Fibers in tangential rows and
in small groups. Prismatic crystals pre-
sent. Cell content in parenchyma cells.
Phellem distinct in polarized light.

Xylem
Rays predominantly uniseriate.

›T

Pith
Pith shape round. Heterogeneous
pith. Thick-walled parenchyma cells
present. Cells dimorphic. Cell content
present (dark staining substances). Pits
in transverse and in longitudinal cell
walls. Vascular bundles not distinct.
Tracheary elements of metaxylem in
Cistus ladanifer L.

distinct radial rows. Axial cells in regu-


lar rows (radial section).

›T

230
Stem xylem: 1 4 9 13 21 22 24 30 40.1 50.1 52.3 62 70 76 78 96 97 109 116.1 136 136.1 141.1
Twig bark: B1 B4 B6 B12 B14 B20 B28 B31 B33
Pith: P1 P3 P3.3 P4 P5 P9 P9.1 P10.2 P12 P13

Cistaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted latewood fibers and by
marginal parenchyma bands. Wood
semi-ring-porous. Vessels predomi-
nantly solitary. Mean tangential diam-
eter of earlywood vessels less than 20
μm, 100-200 vessels per mm2. Fibers
very thick-walled. Axial parenchyma
diffuse and scanty paratracheal. Rays
12-20 per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits opposite and alternate, minute
(less than 4 μm in diameter). Vessel-
ray pits with distinct borders, similar
to intervessel pits in size and shape
throughout the ray cell. Earlywood ves-
sel element length 100-200 μm. Fibers
with distinctly bordered pits (fiber tra-
cheids). Rays with procumbent, square
and upright cells mixed throughout the
ray. Prismatic crystals in rays and in
axial parenchyma cells.

›T ›T

Tangential section
Rays uniseriate and 2 to 3 cells wide.
Cistus ladanifer L.

Wood basic density: 0,65 g/cm³

›T ›T

231
Cistus monspeliensis L.
Narrow-Leaved Cistus
Cistaceae

PLANT DESCRIPTION
Shrub up to 1 m high. Leaves linear-
lanceolate. It occurs in the Mediter-
ranean countries and the Atlantic
islands. In Cyprus it grows on rocky
slopes, in forests and in shrublands
(0-600 m alt.) on the western part of
the island. Cyprus is the easternmost
boundary of the species distribution.

TWIG ANATOMY
Bark
Groups of sieve tube present. Fibers in
tangential rows. Prismatic crystals and
crystal druses present. Phellem homo-
geneous and distinct in polarized light.
All parenchyma cells filled with dark
stained substances.

Xylem
Wood diffuse-porous. No tyloses pre-
sent. Rays exclusively uniseriate. All
ray cells upright and square. More than
20 rays per mm. The first ring consist
of few vessels and very thick-walled
fibers.
›T

Pith
Pith shape oval. Medullary sheath
present. Cells dimorphic. Crystal
druses present. Pits in transverse and
in longitudinal cell walls. Pits grouped
Cistus monspeliensis L.

in transverse walls. Pits of two distinct


size. Vascular bundles not distinct.

›T ›T

232
Stem xylem: 1 4 5 9 13 22 24 30 36 40.2 50.1 53.1 56 58 61 70 76 78 96.1 97 106 116.1
Twig bark: B1 B9 B12 B20 B22 B31 B33
Pith: P1 P2 P4.2 P6.2 P9 P9.1 P9.2 P9.3 P10.2

Cistaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by a
lack of vessels in terminal latewood.
Wood semi-ring-porous to diffuse-
porous. Vessels predominantly solitary.
Mean tangential diameter of earlywood
vessels 20-50 μm, 100-200 vessels per
mm2. Tyloses with thin walls common.
Gums and other deposits in heartwood
vessels and fibers. Fibers very thick-
walled. Axial parenchyma diffuse. Axial
parenchyma scanty paratracheal in
earlywood.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, minute (less than 4 μm
in diameter). Vessel-ray pits with dis-
tinct borders, similar to intervessel pits
in size and shape throughout the ray
cell. Helical thickenings in vessel ele-
ments present. Earlywood vessel ele-
ment length 200-500 μm. Fibers with
simple to minutely bordered pits (libri-
form fibers). Body ray cells procumbent
with one row of upright and square
marginal cells.

›T ›T

Tangential section
Rays uniseriate to 3 cells wide.
Cistus monspeliensis L.

Wood basic density: 0,78 g/cm³

›T ›T

233
Cistus parvifolius Lam.
Rockrose
Cistaceae

PLANT DESCRIPTION
Shrub up to 1 m high. Leaves glau-
cous-green, distinctly three-nerved.
It occurs in the central and eastern
Mediterranean countries. A common
indigenous species to Cyprus growing
on rocky slopes, in forests and thickets
(usually 0-300 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube in tangential
rows. Distinct parenchyma cells dila-
tation. Fibers in tangential rows and
grouped. Prismatic crystals present.
Cortex consist of large and small
thin-walled parenchyma cells. Phellem
distinct in polarized light.

Xylem
As stem wood.

›T

Pith
Pith shape round. Heterogeneous pith.
Thick-walled parenchyma cells present.
Cells dimorphic. Crystal druses pre-
sent. Pits in transverse and in longitu-
Cistus parvifolius Lam.

dinal cell walls. Vascular bundles not


distinct. Tracheary elements of metax-
ylem in distinct radial rows. Axial cells
in regular rows (radial section).

›T ›T

234
Stem xylem: 1 5 9 13 22 25 30 40.1 50.2 53.1 56 58 62 70 76 78 86 96.1 97 103 105 116.2
Twig bark: B2 B3.2 B9 B12 B14 B20 B31 B33
Pith: P1 P3.3 P4 P6.2 P9 P9.1 P10.2 P12 P13

Cistaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by a
lack of vessels in terminal latewood
and by thick-walled latewood fibers.
Wood diffuse-porous. Vessels predomi-
nantly solitary. Mean tangential diam-
eter of earlywood vessels less than 20
μm. More than 200 vessels per mm2.
Tyloses with thin walls common. Gums
and other deposits in heartwood ves-
sels. Fibers very thick-walled. Axial
parenchyma diffuse and scanty para-
tracheal. Apotracheal parenchyma in
narrow bands or lines up to three cells
wide. More than 20 rays per mm.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in diame-
ter). Vessel-ray pits with distinct bor-
ders, similar to intervessel pits in size
and shape throughout the ray cell.
Earlywood vessel element length 200-
500 μm. Fibers with distinctly bordered
pits (fiber tracheids). All ray cells up-
right and square.

›T ›T

Tangential section
Rays of two distinct sizes: predomi-
nantly uniseriate, rarely more than 4
seriate.
Cistus parvifolius Lam.

Wood basic density: 0,79 g/cm³

›T ›T

235
Cistus salvifolius L.
Sage-Leaved Rockrose
Cistaceae

PLANT DESCRIPTION
Shrub up to 1 m high with smooth,
hairy stems. Leaves opposite, ovate
or oblong, with upper surface rugose,
lower surface thinly pilose, and dis-
tinctly veined. Widespread in the Medi-
terranean. Indigenous and common on
Cyprus, growing in garigue, maquis
and pine forests (0-1400 m alt.).

TWIG ANATOMY
Bark
Sclerenchyma cells in phloem and in
cortex. Fibers in tangential rows and
grouped. Sclereids in groups. Prismatic
crystals and crystal druses present.
Phellem homogeneous, distinct in po-
larized light.

Xylem
Vessels predominantly solitary.

›T

Pith
Pith shape round. Heterogeneous pith.
Thick-walled parenchyma cells present.
Cells dimorphic. Cell content present
(dark staining substances). Crystal
druses present. Pits in transverse and
in longitudinal cell walls, grouped. Vas-
Cistus salvifolius L.

›T
cular bundles clearly separate to not
distinct. Tracheary elements of metax-
ylem in distinct radial rows. Axial cells
in regular rows (radial section).

›T

236
Stem xylem: 1 4 9 9.1 13 22 25 30 31 40.2 50.1 53.1 58 60 61 69 70.2 75 96 106 109 116.2 136 136.1 144 145
Twig bark: B7 B9 B12 B14 B15 B19 B20 B22 B31 B33
Pith: P1 P3.3 P4 P5 P6.2 P9 P9.1 P9.2 P10.1 P10.2 P12 P13

Cistaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood semi-ring-porous. Vessels
solitary and in radial multiples of 2 to
4. Mean tangential diameter of early-
wood vessels 20-50 μm, 100-200 ves-
sels per mm2. Gums and other deposits
in heartwood vessels. Fibers thin- to
thick-walled. Tension wood present.
Axial parenchyma absent or extremely
rare or not to recognizable. More than
20 rays per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
In earlywood vessel-ray pits rounded
or angular with large apertures. Ear-
lywood vessels length 200-500 μm.
Vascular and/or vasicentric tracheids
present. Fibers with simple to minutely
bordered pits (libriform fibers). Body
ray cells procumbent with one row of
upright and square marginal cells. Rays
with procumbent, square and upright
cells mixed throughout the ray. Pris-
›T ›T matic crystals and crystal druses in ray
parenchyma cells.
Tangential section
Rays exclusively uniseriate.
Cistus salvifolius L.

Wood basic density: 0,68 g/cm³

›T ›T

237
Fumana arabica (L.) Spach.
Arabian Fumana
Cistaceae

PLANT DESCRIPTION
Subshrub up to 60 cm high. Stems
usually wiry, spreading, suberect,
densely hairy. Leaves alternate, linear.
It occurs in countries of the central and
eastern Mediterranean and eastward
to Iran. Common species in Cyprus,
growing on rocky slopes, maquis and
garigue (0-1200 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and sieve tubes
present. Fibers in tangential rows, scat-
tered or irregularly dispersed. Phellem
homogeneous, distinct in polarized
light.

Xylem
Axial parenchyma diffuse. The first
ring consist of few small vessels and
very thick fibers.

›T

Pith
Pith shape round. Medullary sheath
Fumana arabica (L.) Spach.

present. Heterogeneous pith. Thick-


walled parenchyma cells present. Cells
dimorphic. Cell content present (dark
staining substances). Pits in transverse
and in longitudinal cell walls. Vascular
bundles not distinct. Tracheary ele-
ments of metaxylem in distinct radial
rows. Axial cells in regular rows (radial
section).

›T ›T

238
Stem xylem: 1 5 9 13 21 22 24 30 39.1 40.1 50.1 52.3 62 70 78 96 105 116.2
Twig bark: B1 B4 B12 B13 B31 B33
Pith: P1 P2 P3.3 P4.2 P5 P9 P9.1 P10.2 P12 P13

Cistaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers and by a lack of vessels in ter-
minal latewood. Wood diffuse-porous,
vessels predominantly solitary. Vessels
cell wall thick (more than 2 μm). Mean
tangential diameter of earlywood ves-
sels less than 20 μm, 100-200 vessels
per mm2. Fibers very thick-walled. Axial
parenchyma scanty paratracheal. More
than 20 rays per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits opposite and alternate, minute
(less than 4 μm in diameter). Vessel-
ray pits with distinct borders, similar
to intervessel pits in size and shape
throughout the ray cell. Earlywood ves-
sel element length 100-200 μm. Fib-
ers with distinctly bordered pits (fiber
tracheids). All ray cells upright.

›T ›T

Tangential section
Rays exclusively uniseriate.
Fumana arabica (L.) Spach.

Wood basic density: 0,80 g/cm³

›T ›T

239
Fumana thymifolia (L.) Webb
Thyme-Leaved Fumana
Cistaceae

PLANT DESCRIPTION
Subshrub up to 25 cm high. Young
stems thinly or densely hairy. Leaves
opposite or alternate, lanceolate or
narrowly obovate. It occurs in coun-
tries of the Mediterranean and east-
ward to Iran. Common species on
Cyprus, growing on rocky slopes, sand
dunes and garigue (0-1000 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube present. Few scle-
reids in tangential rows or irregularly
dispersed. Prismatic crystals present.
Cell content in parenchyma cells. Cor-
tex consist of two distinct layers: one
with thin-walled unlignified cells, the
second with lignified cells.

Xylem
The first ring consist of few small ves-
sels and very thick fibers.

›T

Pith
Fumana thymifolia (L.) Webb

Pith shape round to polygonal. Medul-


lary sheath present. Heterogeneous
pith. Thick-walled parenchyma cells
present. Cell content present (dark
staining substances). Pits in transverse
and in longitudinal cell walls. Vascular
bundles not distinct.

›T ›T

240
Stem xylem: 1 4 9 11 13 21 22 24 30 40.1 50.1 52.2 58 62 70 76 78 89 89.1 96 99.2 105 116.2
Twig bark: B1 B17 B18 B20 B28 B35
Pith: P1 P2 P3.3 P5 P9 P9.1 P10.2

Cistaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers and by a lack of vessels in termi-
nal latewood. Wood semi-ring-porous.
Vessels solitary and in clusters. Mean
tangential diameter of earlywood ves-
sels less than 20 μm 100-200 vessels
per mm2. Gums and other deposits in
heartwood vessels and in ray parenchy-
ma cells. Fibers very thick-walled. Axial
parenchyma diffuse and scanty paratra-
cheal. Apotracheal parenchyma in mar-
ginal or in seemingly marginal bands.
Parenchyma marginal thin-walled. Stem
›T ›T
lobed. More than 20 rays per mm.

Radial section
Simple perforation plates. Inter-vessel
pits opposite and alternate, minute
(less than 4 μm in diameter). Vessel-
ray pits with distinct borders, similar
to intervessel pits in size and shape
throughout the ray cell. Earlywood ves-
sel element length 50-100 μm. Fibers
with distinctly bordered pits (fiber trac-
heids). All ray cells upright and square.

›T ›T

Tangential section Fumana thymifolia (L.) Webb


Rays exclusively uniseriate.

Wood basic density: 0,82 g/cm³

›T ›T

241
Helianthemum obtusifolium Dunal
Cistaceae

PLANT DESCRIPTION
Subshrub up to 25 cm high, with
densely tomentose, suberect stems.
Leaves opposite, lamina lanceolate or
oblong-elliptic. Endemic to Cyprus, it
grows on rocky slopes, in maquis and
garigue (0-900 m alt.).

TWIG ANATOMY
Bark
No slide available.

Xylem
The first ring consist of few small ves-
sels and very thick fibers.
Helianthemum obtusifolium Dunal

Pith
Pith not visible in polarized light. Pith
shape round to oval. Medullary sheath
present. Cell content present (dark
staining substances). Vascular bundles
not distinct. Tracheary elements of
metaxylem in distinct radial rows.

›T

242
Stem xylem: 1 5 9 13 21 22 24 30 40.1 50.1 52.3 58 62 69 70 76 78 89 89.1 96 99.2 105 116.2
Twig bark: No slide available.
Pith: P0.1 P1 P2 P5 P10.2 P12

Cistaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted and thick-walled late-
wood fibers. Wood diffuse-porous.
Vessels predominantly solitary. Mean
tangential diameter of earlywood ves-
sels less than 20 μm, 100-200 vessels
per mm2. Gums and other deposits in
heartwood vessels. Fibers thick- to very
thick-walled. Axial parenchyma diffuse
and scanty paratracheal. Apotracheal
parenchyma in marginal or in seem-
ingly marginal bands, dark in polarized
light. Stem lobed. More than 20 rays
per mm.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits opposite and alternate, minute
(less than 4 μm in diameter). Vessel-
ray pits with distinct borders, similar
to intervessel pits in size and shape
throughout the ray cell. Earlywood ves-
sel element length 100-200 μm. Fib-
ers with distinctly bordered pits (fiber
tracheids). All ray cells upright.

Helianthemum obtusifolium Dunal


›T ›T

Tangential section
Rays exclusively uniseriate.

Wood basic density: 0,71 g/cm³

›T ›T

243
Helianthemum stipulatum (Forssk.) C.Chr.
Cistaceae

PLANT DESCRIPTION
Subshrub up to 40 cm high, with
densely tomentose stems. Leaves op-
posite, ovate or almost linear, with up-
per surface thinly hairy, lower surface
densely grey-tomentose. In Cyprus it
grows on sandy beaches and rocky
slopes (0-200 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclereids in
tangential rows and scattered or ir-
regularly dispersed. Prismatic crystals
present. Cell content in parenchyma
cells. Cortex consist of large and small
thin-walled parenchyma cells. Phellem
distinct in polarized light.
Helianthemum stipulatum (Forssk.) C.Chr.

Xylem
Rays predominantly uniseriate. The
first ring consist of few small vessels
and very thick fibers.
›T

Pith
Pith shape round. Medullary sheath
present. Heterogeneous pith. Thick-
walled parenchyma cells present.
Cell content present (dark staining
substances). Pits in transverse cell
walls. Vascular bundles not distinct.
Tracheary elements of metaxylem in
distinct radial rows.

›T

244
Stem xylem: 1 4 5 9 13 21 22 24 30 40.1 50.1 52.2 56 58 60 62 69 70 76 78 89 96 97 105 116.2 136 136.1
Twig bark: B1 B4 B17 B18 B20 B28 B31 B33
Pith: P1 P2 P3.3 P5 P9.1 P10.2 P12

Cistaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers and by a lack of vessels in termi-
nal latewood. Wood semi-ring-porous
to diffuse-porous. Vessels predomi-
nantly solitary. Mean tangential diam-
eter of earlywood vessels less than 20
μm, 100-200 vessels per mm2. Tyloses
with thin walls common. Gums and
other deposits in heartwood vessels.
Fibers thick- to very thick-walled. Axial
parenchyma diffuse, scanty paratra-
cheal and in marginal or in seemingly
marginal bands. More than 20 rays per
›T ›T
mm.

Radial section
Simple perforation plates. Inter-vessel
pits opposite and alternate, minute
(less than 4 μm in diameter). Vessel-
ray pits with distinct borders, similar
to intervessel pits in size and shape
throughout the ray cell. Earlywood
vessel element length 50-100 μm.
Vascular and/or vasicentric tracheids
present. Fibers with distinctly bordered
pits (fiber tracheids). All ray cells up-

Helianthemum stipulatum (Forssk.) C.Chr.


right and square. Prismatic crystals in
ray parenchyma cells.

›T ›T

Tangential section
Rays uniseriate to 3 cells wide.

Wood basic density: 0,60 g/cm³

›T ›T

245
Helianthemum syriacum (Jacq.) Dum-Cours.
Cistaceae

PLANT DESCRIPTION
Subshrub up to 40 cm high, with
densely tomentose stems. Leaves op-
posite, lanceolate or narrowly ovate,
densely grey-tomentose above and
below. Indigenous in the Mediterranean
countries. In Cyprus it grows on sandy
beaches and rocky slopes with garigue
and phrygana vegetation (0-800 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube present. Some
rays become dilated. Sclerenchyma
cells both in phloem and in cortex. Fib-
ers scattered or irregularly dispersed.
Prismatic crystals and crystal druses
present. Cell content in parenchyma
cells. Phellem distinct in polarized
light.
Helianthemum syriacum (Jacq.) Dum-Cours.

Xylem
The first ring consist of few small ves-
sels and very thick fibers.

›T

Pith
Pith not visible in polarized light. Pith
shape round. Medullary sheath pre-
sent. Cell content present (dark stain-
ing substances). With prismatic crystals
and crystal druses. Vascular bundles
clearly separate to not distinct. Tra-
cheary elements of metaxylem in dis-
tinct radial rows.

›T ›T

246
Stem xylem: 1 5 9 9.1 13 21 22 24 30 40.1 50.1 53.1 62 69 70 78 96.1 97 105 116.2
Twig bark: B1 B6 B7 B13 B20 B22 B28 B33
Pith: P0.1 P1 P2 P5 P6 P6.2 P10.1 P10.2 P12

Cistaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood diffuse-porous. Vessels
solitary or short radial multiples. Mean
tangential diameter of earlywood ves-
sels less than 20 μm, 100-200 vessels
per mm2. Fibers thick- to very thick-
walled. Axial parenchyma scanty para-
tracheal. More than 20 rays per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits opposite and alternate, minute
(less than 4 μm in diameter). Vessel-
ray pits with distinct borders, similar
to intervessel pits in size and shape
throughout the ray cell. Earlywood ves-
sel element length 200-500 μm. Fib-
ers with distinctly bordered pits (fiber
tracheids). All ray cells upright and

Helianthemum syriacum (Jacq.) Dum-Cours.


square.

›T ›T

Tangential section
Rays uniseriate and up to 3 cells wide.

Wood basic density: 0,57 g/cm³

›T ›T

247
Convolvulus dorycnium L.
Bindweed
Convolvulaceae

PLANT DESCRIPTION
Erect, much branched subshrub 30-80
cm high, with distinctly hairy stems,
especially towards the base. Leaves al-
ternate, linear. An east Mediterranean
species, extending eastward to Iran. In
Cyprus it occurs on dry stony places,
alongroad sides and fallow land (0-300
m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube present.
Collapsed sieve tubes present. Some
rays become dilated. Sclereids scat-
tered or irregularly dispersed. Crystal
druses present. Phellem homogene-
ous.

Xylem
Vessels predominantly solitary. Early-
wood vessel element length 200-500
μm. Axial parenchyma rare, scanty
paratracheal. All ray cells upright and
squared. More than 20 rays per mm.
›T

Pith
Pith not visible in polarized light. Pith
shape polygonal. Medullary sheath
Convolvulus dorycnium L.

present. Heterogenous pith. Fibers pre-


sent. Unlignified cells present. Prismat-
ic crystals present. Pits in transverse
and in longitudinal cell walls. Vascular
bundles in the pith. Vascular bundles
not distinct.

›T ›T

248
Stem xylem: 1 4 11 13 22 26 30 31 39.1 41 45 52.3 62 78 79 89 89.1 96 97 100.2 102 103 109 116.1 144 145
Twig bark: B1 B4 B6 B18 B22 B31
Pith: P0.1 P1.1 P2 P3 P3.2 P3.4 P6 P9 P9.1 P10 P10.2

Convolvulaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
large marginal parenchyma bands.
Wood semi-ring-porous. Vessel clusters
common. Vessels cell wall thick (more
than 2 μm). Mean tangential diameter
of earlywood vessel lumina 50-100 μm.
Vessels of two distinct diameter class-
es. Axial parenchyma scanty paratra-
cheal to vasicentric. Axial parenchyma
in marginal or in seemingly marginal
bands. Parenchyma marginal thin-
walled, dark in polarized light. Rays
12-20 per mm.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, medium (7-10 μm in
diameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Vessel-ray pits rounded or angular with
large apertures. Earlywood vessel ele-
ment length 100-200 μm. Fibers with
distinctly bordered pits (fiber trac-
heids). Rays with procumbent, square
and upright cells mixed throughout the
ray. Crystal druses in ray parenchyma
cells.

›T ›T

Tangential section
Rays of two distinct sizes: uniseriate
and 4 to 5 seriate. Larger rays height
Convolvulus dorycnium L.

more than 1 mm. Rays disappear in


polarized light.

Wood basic density: 0,29 g/cm³

›T ›T

249
Convolvulus oleifolius var. desertii Desr. Pamp.
Convolvulaceae

PLANT DESCRIPTION
Subshrub up to 50 cm high, with erect
or spreading, densely hairy stems,
branched from the base. Leaves alter-
nate, narrowly linear. It grows in Libya
and Cyprus, restricted to dry, rocky,
calcareous slopes with phrygana veg-
etation (150-200 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube present. Groups
of sieve tube in radial rows. Collapsed
sieve tubes present. Fibers with unlig-
nified innermost layer present. Crystal
druses present. With secretory ele-
ments in ducts. Cortex consist of large
Convolvulus oleifolius var. desertii Desr. Pamp.

and small thin-walled parenchyma


cells. Phellem homogeneous.

Xylem
Mean tangential diameter of earlywood
vessels lumina less than 20 μm. Rays
uniseriate. More than 20 rays per mm.
›T

Pith
Pith not visible in polarized light. Pith
shape round. Heterogeneous pith.
Unlignified cells present. Cells dimor-
phic. Crystal druses present. Vascular
bundles in the pith. Vascular bundles
clearly separate. Tracheary elements of
metaxylem in distinct radial rows.

›T ›T

250
Stem xylem: 1 5 9 13 22 26 30 40.2 45 50.1 53.1 61 69 78 89 89.1 96.1 97 105 116.2
Twig bark: B1 B3 B4 B22 B25 B26 B31
Pith: P0.1 P1 P3.4 P4.1 P6.2 P10 P10.1 P12

Convolvulaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands. Wood
diffuse-porous. Vessels predominantly
solitary. Mean tangential diameter of
earlywood vessels 20-50 μm, 100-200
vessels per mm2. Vessels of two dis-
tinct diameter classes. Fibers thin- to
thick-walled. Axial parenchyma scanty
paratracheal. Apotracheal parenchyma
in marginal or in seemingly marginal
bands. Parenchyma marginal thin-
walled, dark in polarized light. En-
closed sieve tubes. More than 20 rays
per mm.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, medium (7-10 μm in
diameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 200-
500 μm. Fibers with simple to minutely

Convolvulus oleifolius var. desertii Desr. Pamp.


bordered pits (libriform fibers). All ray
cells upright and square.

›T ›T

Tangential section
Rays predominantly uniseriate.

Wood basic density: 0,60 g/cm³

›T ›T

251
Convolvulus oleifolius Desr. var. oleifolius
Convolvulaceae

PLANT DESCRIPTION
Subshrub up to 50 cm high, with erect
or spreading, densely hairy stems,
branched from the base. Leaves alter-
nate, oblanceolate. Indigenous to cen-
tral and east Mediterranean countries.
In Cyprus it grows in garigue and on
dry, rocky slopes (0-700 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclereids scattered
or irregularly dispersed. Fibers with
unlignified innermost layer present.
Crystal druses present. Cortex consist
of large and small thin-walled paren-
chyma cells. Phellem homogeneous.
Epidermis distinct in polarized light.
Convolvulus oleifolius Desr. var. oleifolius

Xylem
Thick-walled fibers. Axial parenchyma
absent or extremely rare or not to rec-
ognisable.
›T

Pith
Pith not visible in polarized light. Pith
shape round. Cells dimorphic. Crystal
druses present. Pits in transverse and
in longitudinal cell walls. Vascular bun-
dles not distinct. Tracheary elements
of metaxylem in distinct radial rows.

›T ›T

252
Stem xylem: 1 5 9 11 13 22 25 30 39.1 40.2 45 50.1 53.1 62 69 78 96.1 105 116.2 144 146
Twig bark: B1 B4 B9 B10 B18 B22 B31 B33
Pith: P0.1 P1 P4.1 P6.2 P9 P9.1 P10.2 P12

Convolvulaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
thick-walled latewood fibers and by a
lack of vessels in terminal latewood.
Wood diffuse-porous. Vessels solitary
or in clusters. Vessels cell wall thick
(more than 2 μm). Mean tangential
diameter of earlywood vessels 20-50
μm, 100-200 vessels per mm2. Vessels
of two distinct diameter classes. Fibers
thin- to thick-walled. Axial parenchyma
scanty paratracheal. More than 20 rays
per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 200-
500 μm. Fibers with distinctly bor-
dered pits (fiber tracheids). All ray cells
upright and square. Crystal druses in
axial parenchyma cells.

Convolvulus oleifolius Desr. var. oleifolius


›T ›T

Tangential section
Rays predominantly uniseriate.

Wood basic density: 0,67 g/cm³

›T ›T

253
Convolvulus oleifolius Desr. var. pumilus Pamp.
Convolvulaceae

PLANT DESCRIPTION
Much branched subshrub up to 15 cm
high, forming dense, rounded tufts.
Leaves alternate, oblanceolate, densely
covered with silvery hairs. In Cyprus it
is restricted to the southern costal area
(0-20 m alt.) where it grows on rocky
ground or among low shrubs. Found
also in Libya.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Some rays become
dilated. Crystal druses present. Secre-
tory elements in ducts present. Cortex
consist of large and small thin-walled
parenchyma cells. Phellem homogene-
Convolvulus oleifolius Desr. var. pumilus Pamp.

ous.

Xylem
Mean tangential diameter of earlywood
vessels lumina less than 20 μm. Rays
exclusively uniseriate. Ray dimorphism
absents.
›T

Pith
Pith not visible in polarized light. Pith
shape round. Medullary sheath pre-
sent. Heterogeneous pith. Unlignified
cells present. Cells dimorphic. Vascu-
lar bundles in the pith. Vascular bun-
dles not distinct. Tracheary elements
of metaxylem in distinct radial rows.
Axial cells in regular rows (radial sec-
tion).

›T ›T

254
Stem xylem: 1 3 9 9.1 13 21 22 25 39.1 40.2 49 53.1 62 69 70 78 89 89.1 96 97 100.2 102 103 105 116.2 144 145
Twig bark: B1 B4 B6 B22 B25 B26 B31
Pith: P0.1 P1 P2 P3.4 P4 P10 P10.2 P12 P13

Convolvulaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands. Wood
ring-porous. Vessels solitary and in ra-
dial multiples of 2 to 4 common. Ves-
sels cell wall thick (more than 2 μm).
Mean tangential diameter of earlywood
vessels 20-50 μm, 40-100 vessels per
mm2. Fibers thick- to very thick-walled.
Axial parenchyma scanty paratracheal
and in marginal or in seemingly mar-
ginal bands. Parenchyma marginal
thin-walled, dark in polarized light.
More than 20 rays per mm.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits opposite and alternate, small (4-7
μm in diameter). Earlywood vessel ele-
ment length 200-500 μm. Fibers with
distinctly bordered pits (fiber trac-
heids). All ray cells upright and square.
Crystal druses in ray parenchyma cells.

Convolvulus oleifolius Desr. var. pumilus Pamp.


›T ›T

Tangential section
Rays of two distinct sizes: uniseriate
and 2-3 cells wide. Larger rays height
more than 1 mm. Rays disappear in
polarized light.

Wood basic density: 0,83 g/cm³

›T ›T

255
Cornus sanguinea L.
Common Dogwood
Cornaceae

PLANT DESCRIPTION
Deciduous shrub, growing 2–6 m high,
with dark greenish-brown branches
and twigs. Leaves opposite, ovate to
oblong with entire margin, deeply
veined. Its natural range covers most
of Europe and western Asia. It is espe-
cially abundant in riversides, especially
in shady areas and ravines.

TWIG ANATOMY
Bark
Sclerenchyma cells in phloem and in
cortex. Fibers scattered or irregularly
dispersed and in groups. Prismatic
crystals and druses present. Cell con-
tent in parenchyma cells. Cortex
homogeneous. Epidermis distinct in
polarized light.

Xylem
Mean tangential diameter of earlywood
vessel lumina 20-50 μm. Rays exclu-
sively uniseriate.

›T

Pith
Pith not visible in polarized light. Pith
shape round. Medullary sheath pre-
sent. Heterogeneous pith. Unlignified
cells present. With crystal druses and
crystal sand present. Tracheary ele-
ments of metaxylem in distinct radial
Cornus sanguinea L.

rows.

›T

256
Stem xylem: 1 5 9 14 17 21 26 31 41 49 53.1 62 70 77 97 107 116.2
Twig bark: B7 B9 B13 B14 B20 B22 B28 B31 B14.3
Pith: P0.1 P1 P2 P3 P3.4 P6.2 P6.4 P12

Cornaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood diffuse-porous. Vessels
predominantly solitary. Mean tan-
gential diameter of earlywood vessel
lumina 50-100 μm, 40-100 vessels per
mm2. Fibers very thick-walled. Axial pa-
renchyma diffuse in aggregates. More
than 20 rays per mm.

›T ›T

Radial section
Scalariform perforation plates with
20-40 bars. Inter-vessel pits opposite,
medium (7-10 μm in diameter). Vessel-
ray pits rounded or angular with large
apertures. Earlywood vessel element
length 200-500 μm. Fibers with dis-
tinctly bordered pits (fiber tracheids).
Body ray cells procumbent with mostly
2-4 rows of square marginal cells.

›T ›T

Tangential section
Ray width predominantly 1 to 3 cells.
Cornus sanguinea L.

Wood basic density: 0,61 g/cm³

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257
Lomelosia cyprica (Post) Greuter et Burdet
= Scabiosa cyprica Post
Dipsacaceae

PLANT DESCRIPTION
Shrub up to 1,2 m high. Leaves oppo-
site, hairy and greyish-green. An en-
demic species, occurring on dry slopes
in garigue and open pine forests on
both igneous and limestone forma-
tions (150-1200 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Some rays become
dilated. Prismatic crystals present.
Phellem consist of large thin-walled
cork cells.

Xylem
Mean tangential diameter of earlywood
vessels lumina less than 20 μm. Fibers
Lomelosia cyprica (Post) Greuter et Burdet

very thick-walled.

›T

Pith
Pith shape round. Cells dimorphic. Pits
in transverse and in longitudinal cell
walls. Pits grouped. Pits of two distinct
size. Vascular bundles clearly separate
to not distinct. Tracheary elements of
metaxylem in distinct radial rows.

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258
Stem xylem: 1 4 9 13 22 25 30 40.2 50.1 53.1 62 69 70.3 75 76 96 105 116.2
Twig bark: B1 B4 B6 B20 B35
Pith: P1 P4 P9 P9.1 P9.2 P9.3 P10.1 P10.2 P12

Dipsacaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted and thick-walled late-
wood fibers and by a lack of vessels
in terminal latewood. Wood semi-ring-
porous. Vessels predominantly solitary.
Mean tangential diameter of earlywood
vessels 20-50 μm, 100-200 vessels
per mm2. Fibers thin- to thick-walled.
Radial flat marginal fibers. Axial paren-
chyma extremely rare, diffuse. More
than 20 rays per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in diame-
ter). Vessel-ray pits with distinct bor-
ders, similar to intervessel pits in size
and shape throughout the ray cell. Ear-
lywood vessel element length 200-500
μm. Fibers with distinctly bordered pits
(fiber tracheids). All ray cells upright
and square.

Lomelosia cyprica (Post) Greuter et Burdet


›T ›T

Tangential section
Rays exclusively uniseriate.

Wood basic density: 0,53 g/cm³

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259
Pterocephalus multiflorus Poech subsp. multiflorus
Pterocephalus
Dipsacaceae

PLANT DESCRIPTION
Much branched shrub up to 1 m high,
with erect or spreading branches.
Leaves opposite, elliptic, tapering at
apex and base. Endemic to Cyprus. It
occurs in phrygana, on rocky slopes,
maquis and open pine forests on lime-
stone and igneous formations (150-
1900 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube in radial rows.
Prismatic crystals and acicular present.
Phellem consist of large thin-walled
Pterocephalus multiflorus Poech subsp. multiflorus

cork cells.

Xylem
As stem wood.

›T

Pith
Pith round in shape. Cells dimorphic.
Crystal druses present. Pits transverse
and in longitudinal cell walls (radial
section). Vascular bundles not distinct.
Axial cells in regular rows (radial sec-
tion).
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260
Stem xylem: 1 4 7 11 13 22 25 30 31 36 40.2 45 49 53.1 62 69 76 78 96.1 105 116.2
Twig bark: B1 B3 B20 B21
Pith: P1 P4.1 P6.2 P9 P9.1 P10.2 P13

Dipsacaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers and by a lack of vessels in termi-
nal latewood. Wood semi-ring-porous.
Vessels in diagonal and radial pattern,
predominantly in clusters. Mean tan-
gential diameter of earlywood vessels
20-50 μm, 40-100 vessels per mm2.
Vessels of two distinct diameter class-
es. Fibers thin- to thick-walled. Axial
parenchyma diffuse and scanty paratra-
cheal. More than 20 rays per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.

Pterocephalus multiflorus Poech subsp. multiflorus


Vessel-ray pits rounded or angular with
large apertures. Helical thickenings in
vessel elements present. Earlywood
vessel element length 200-500 μm.
Fibers with distinctly bordered pits
(fiber tracheids). All ray cells upright
and square.

›T ›T

Tangential section
Rays predominantly uniseriate.

Wood basic density: 0,53 g/cm³

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261
Pterocephalus multiflorus Poech subsp. obtusifolius
Holmboe
Dipsacaceae

PLANT DESCRIPTION
Much branched, low shrub up to 1 m
high with erect or spreading branches.
Leaves obovate or subspathulate, ob-
tuse. Endemic to Cyprus. It occurs on
dry calcareous slopes (0-800 m alt.).
Pterocephalus multiflorus Poech subsp. obtusifolius Holmboe

TWIG ANATOMY
Bark
Groups of sieve tube present. Groups
of sieve tube in radial rows. Collapsed
sieve tubes present. Prismatic crystals
present. Phellem consist of large thin-
walled cork cells.

Xylem
As stem wood.

›T

Pith
Pith shape polygonal. Thick-walled
parenchyma cells present. Cells di-
morphic. Crystal druses present. Pits
in transverse and in longitudinal cell
walls. Pits in transverse cell walls. Vas-
cular bundles not distinct.

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262
Stem xylem: 1 3 4 9 11 13 22 24 30 36 40.1 45 50.2 53.2 58 62 64 69 78 96 105 116.2
Twig bark: B1 B3 B4 B20 B30
Pith: P1.1 P3.3 P4 P6.2 P9 P9.1 P10.2

Dipsacaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted and thick-walled late-
wood fibers and by a lack of vessels in
terminal latewood. Wood ring-porous
to semi-ring-porous. Vessels solitary or
in clusters. Mean tangential diameter
of earlywood vessels less than 20 μm.
More than 200 vessels per mm2. Ves-
sels of two distinct diameter classes.
Gums and other deposits in heartwood
vessels. Fibers thin- to thick-walled.
Axial parenchyma scanty paratracheal.
More than 20 rays per mm.
›T ›T

Pterocephalus multiflorus Poech subsp. obtusifolius Holmboe


Radial section
Simple perforation plates. Inter-vessel
pits alternate, minute (less than 4 μm
in diameter). Vessel-ray pits with dis-
tinct borders, similar to intervessel pits
in size and shape throughout the ray
cell. Helical thickenings in vessel ele-
ments present. Earlywood vessel ele-
ment length more than 500 μm. Fibers
with distinctly bordered pits (fiber tra-
cheids). Helical thickenings in ground
tissue fibers. All ray cells upright.

›T ›T

Tangential section
Rays predominantly uniseriate.

Wood basic density: 0,45 g/cm³

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263
Diospiros kaki Thunb.
Japanese Persimmon, Kaki Persimmon
Ebenaceae

PLANT DESCRIPTION
Deciduous tree up to ten m high. Its
leaves are dark green, broadly lanceo-
late, stiff and equally wide as long.
Cultivation of the fruit extended from
East Asia, north and south America and
Europe. Cultivated on Cyprus.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclerenchyma
cells in phloem and in cortex. Fibers
grouped. Sclereids in tangential rows.
Prismatic crystals present. Phellem
homogeneous.

Xylem
Mean tangential diameter of earlywood
vessel lumina 50-100 μm.

›T

Pith
Pith shape round. Thick-walled paren-
chyma cells present. Cells dimorphic.
With prismatic crystals. Pits in trans-
verse and in longitudinal cell walls.
Diospiros kaki Thunb.

Vascular bundles clearly separate to


not distinct. Tracheary elements of
metaxylem in distinct radial rows.

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264
Stem xylem: 1 5 9 9.1 13 21 25 30 39.1 42 45 47 53.1 58 60 61 69 70.2 76 78 86 97 100 108 115 119 120 136 141.1
Twig bark: B1 B4 B7 B9 B14 B15 B17 B20 B31
Pith: P1 P3 P3.3 P4 P6 P9 P9.1 P10.1 P10.2 P12

Ebenaceae
STEM ANATOMY
Transverse section
Growth ring boundaries clearly demar-
cated. Wood diffuse-porous. Vessels
solitary and in radial multiples of 2 to 4
elements. Vessels cell wall thick (more
than 2 μm). Mean tangential diameter
of earlywood vessel lumina 100-200
μm, 5-20 vessels per mm2. Vessels of
two distinct diameter classes. Gums
and other deposits in heartwood ves-
sels. Fibers thin- to thick-walled. Ten-
sion wood present. Paratracheal paren-
chyma scanty. Apotracheal parenchyma
in narrow bands or lines up to three
›T ›T
cells wide. Rays per mm 4-12.

Radial section
Simple perforation plates. Inter-vessel
pits opposite, small (4-7 μm in diam-
eter). Vessels-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessels length 200-500 μm.
Vascular and/or vasicentric tracheids
present. Fibers with simple to minutely
bordered pits (libriform fibers). Body
ray cells procumbent with over 4 rows
of upright and square marginal cells.
Prismatic crystals in axial parenchyma
cells.

›T ›T

Tangential section
Ray width predominantly 1 to 3 cells.
Rays with multiseriate portions as wide
as uniseriate portions. Rays, axial pa-
renchyma and vessel elements storied.
Diospiros kaki Thunb.

Wood basic density: 0,67 g/cm³

›T ›T

265
Elaeagnus angustifolia L.
Oleaster
Elaeagnaceae

PLANT DESCRIPTION
Deciduous shrub or small tree up to 8
m high; leaves with silvery, flat, scaly
hairs. A naturalized species in Cyprus,
naturalized on uncultivated slopes and
cultivated in gardens, parks and road-
side plantations as ornamental (0-1400
m alt.). Native to western Asia extending
eastward up to northern China.

TWIG ANATOMY
Bark
Groups of sieve tube present. Groups
of sieve tube in tangential rows.
Collapsed sieve tubes present. Scle-
reids in tangential rows and scattered
or irregularly dispersed. Crystal sand
present. Cell content in parenchyma
cells. Phellem homogeneous. Epider-
mis distinct in polarized light.

Xylem
Mean tangential diameter of earlywood
vessels lumina 20-50 μm. Rays pre-
dominantly uniseriate. More than 20
rays per mm.
›T

Pith
Pith shape polygonal. Cells dimorphic.
Cell content present (dark staining
Elaeagnus angustifolia L.

substances). Crystal sand present. Pits


in transverse and in longitudinal cell
walls. Pits grouped. Vascular bundles
not distinct.

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266
Stem xylem: 1 3 7 9 9.1 11 13 22 25 30 36 41 50.1 52.3 60 62 69 70.2 76 78 86 96 98 103 104 106 114
Twig bark: B1 B2 B4 B17 B18 B23 B28 B31 B33
Pith: P1.1 P4 P5 P6.4 P9 P9.1 P9.2 P10.2

Elaeagnaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
the difference in vessel size between
latewood and earlywood, and by zones
with fibers of variable cell wall thick-
ness. Wood ring-porous. Vessels in
diagonal pattern, solitary, in clusters
or in radial multiples of 2 to 4. Mean
tangential diameter of earlywood ves-
sel lumina 50-100 μm, 100-200 vessels
per mm2. Fibers thin- to thick-walled.
Tension wood present. Axial parenchy-
ma diffuse. Axial parenchyma scanty
paratracheal. Apotracheal parenchyma
›T ›T
in narrow bands or lines up to three
cells wide. Less than 4 rays per mm.

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Helical thickenings in vessel elements
present. Earlywood vessel element
length 100-200 μm. Vascular and/or
vasicentric tracheids present. Fibers
with distinctly bordered pits (fiber tra-
cheids). All ray cells procumbent. Body
ray cells procumbent with one row of
upright and square marginal cells.

›T ›T

Tangential section
Rays of two distinct sizes: uniseriate
and up to 10 seriate.
Elaeagnus angustifolia L.

Wood basic density: 0,54 g/cm³

›T ›T

267
Arbutus andrachne L.
Eastern Strawberry Tree
Ericaceae

PLANT DESCRIPTION
Evergreen shrub, 3-5(10) m high, with
a characteristic smooth bark, which is
reddish in winter peeling off to green
towards summer. Native to Cyprus, com-
mon in medium and high altitude (700-
1500 m). Indigenous to Mediterranean
countries from Albania to Palestine, and
from Crimea to the Caucasus.

TWIG ANATOMY
Bark
Groups of sieve tube in tangential
rows. Collapsed sieve tubes present.
Some rays become dilated. Sclereids
scattered or irregularly dispersed. Pris-
matic crystals present. Phellem homo-
geneous, distinct in polarized light.

Xylem
Vessels predominately in clusters.
Body ray cells procumbent with over 4
rows of squared marginal cells. More
than 20 rays per mm.

›T

Pith
Pith shape round. Heterogeneous pith.
Thick-walled parenchyma cells present.
Cells dimorphic. Pits in transverse
and in longitudinal cell walls. Vascular
bundles not distinct. Tracheary ele-
Arbutus andrachne L.

ments of metaxylem in distinct radial


rows. Axial cells in regular rows (radial
section).

›T ›T

268
Stem xylem: 1 4 9 11 13 22 25 30 36 40.2 50.2 53.1 61 65 69 76 78 96 97 105 106 107 115
Twig bark: B1 B2 B4 B6 B18 B20 B31 B33
Pith: P1 P3.3 P4 P9 P9.1 P10.2 P12 P13

Ericaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers and by a lack of vessels in termi-
nal latewood. Wood semi-ring-porous.
Vessels solitary or in small clusters.
Mean tangential diameter of earlywood
vessels 20-50 μm, more than 200
vessels per mm2. Fibers thin- to thick-
walled. Axial parenchyma diffuse and
scanty paratracheal. Rays 4-12 per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Helical thickenings in vessel elements
present. Earlywood vessel element
length 200-500 μm. Fibers with simple
to minutely bordered pits (libriform
fibers). Septate fibers present. Ray cells
square, in some ray with 1-3 row of
upright and square marginal cells.

›T ›T

Tangential section
Rays uniseriate to 3 cells wide.
Arbutus andrachne L.

Wood basic density: 0,64 g/cm³

›T ›T

269
Arbutus unedo L.
Strauberry Tree
Ericaceae

PLANT DESCRIPTION
Evergreen shrub, 3-5 m high, with a
characteristic fissured bark. Native to
Cyprus, with a limited distribution,
found only in Akamas peninsula (30-
80 m alt.). It is common in south-east
Europe towards western Ireland and in
the Mediterranean countries as far as
Turkey.

TWIG ANATOMY
Bark
Groups of sieve tube in radial rows.
Prismatic crystals present. Phellem
homogeneous and distinct in polarized
light.

Xylem
Growth ring indistinct or absent. Wood
diffuse-porous. Vessels predominantly
solitary. Some vessel-ray pits with large
horizontal or vertical apertures.

›T

Pith
Pith shape round. Cells dimorphic.
With prismatic crystals in axial rows.
Pits in transverse and in longitudinal
cell walls. Pits of two distinct size in
transverse cell walls. Vascular bundles
not distinct. Axial cells in regular rows
(radial section).
Arbutus unedo L.

›T ›T

270
Stem xylem: 1 4 11 13 22 27 31 36 40.2 50.2 53.1 56 58 61 65 69 70 78 96 97 103 105 106 116.2
Twig bark: B1 B3 B20 B31 B33
Pith: P1 P4.1 P6 P9 P9.1 P9.3 P10.2 P13

Ericaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers and by a lack of vessels in termi-
nal latewood. Wood semi-ring-porous.
Vessel clusters common. Mean tan-
gential diameter of earlywood vessels
20-50 μm, more than 200 vessels per
mm2. Tyloses with thin walls common.
Gums and other deposits in heartwood
vessels. Fibers thick- to very thick-
walled. Axial parenchyma scanty para-
tracheal. More than 20 rays per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, large (more than 10 μm
in diameter). Vessel-ray pits rounded
or angular with large apertures. Helical
thickenings in vessel elements present.
Earlywood vessel element length 200-
500 μm. Fibers with simple to minutely
bordered pits (libriform fibers). Septate
fibers present. All ray cells upright and
square. Body ray cells procumbent with
one row of upright or square marginal
cells.

›T ›T

Tangential section
Rays of two distinct sizes: uniseriate
and 2-3 seriate. Larger rays height
more than 1 mm.
Arbutus unedo L.

Wood basic density: 0,58 g/cm³

›T ›T

271
Calluna vulgaris (L.) Hull
Common Heather
Ericaceae

PLANT DESCRIPTION
A low-growing perennial shrub 20-50
cm high. Small scale-leaves (less than
2–3 mm long) borne in opposite and
decussate pairs. It is found widely in
Europe and Asia Minor on acidic soils
in open sunny situations and in moder-
ate shade.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Phloem uniform.
Layered phellem.

Xylem
As stem wood.

›T

Pith
Pith shape round. Heterogenous pith.
Thick-walled parenchyma cells present.
Calluna vulgaris (L.) Hull

Cells dimorphic. Pits in transverse


and in longitudinal cell walls, in thick-
walled cells, thin-walled cells without
pits. Vascular bundles not distinct.

›T

272
Stem xylem: 1 5 9 13 21 24 30 40.1 50.2 52.3 61 69 76 96 105 115
Twig bark: B1 B4 B8 B31 B29
Pith: P1 P3 P3.3 P4 P9 P9.1 P10.2

Ericaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood diffuse-porous. Vessels
predominantly solitary. Mean tangen-
tial diameter of earlywood vessels less
than 20 μm. More than 200 vessels per
mm2. Fibers thin- to thick-walled. Axial
parenchyma diffuse. Rays 4-12 per
mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits opposite, minute (less than 4 μm
in diameter). Vessel-ray pits with dis-
tinct borders, similar to intervessel pits
in size and shape throughout the ray
cell. Earlywood vessels length 100-200
μm. Fibers with simple to minutely
bordered pits (libriform fibers). All ray
cells upright and square.

›T ›T

Tangential section
Rays uniseriate.
Calluna vulgaris (L.) Hull

Wood basic density: 0,73 g/cm³

›T ›T

273
Erica arborea L.
Tree Heath
Ericaceae

PLANT DESCRIPTION
An evergreen shrub up to 4 m high.
Leaves needle-like. It grows mostly on
acid soil in dry evergreen shrublands,
surrounding the Mediterranean Basin
and west to Portugal, Canary and Ma-
deira islands.

Photo: Piermattei
TWIG ANATOMY
Bark
Groups of sieve tubes and collapsed
sieve tubes present. Some rays become
dilated. Sclerenchyma cells in phloem
and cortex. Fibers and sclereids in
tangential rows. Prismatic crystals pre-
sent. Cell content in parenchyma cells.

Xylem
As stem wood.

›T

Pith
Pith shape round. Cell dimorphic. Cell
content present (dark staining sub-
stances). Pits in transverse and in lon-
gitudinal cell walls. Vascular bundles
clearly separate.
Erica arborea L.

›T ›T

274
Stem xylem: 1 4 9 13 21 22 24 30 40.2 50.1 53.1 61 69 76 77 78 97 98 103 106 107 115
Twig bark: B1 B4 B6 B7 B9 B12 B15 B17 B20 B11
Pith: P1 P4 P5 P9 P9.1 P10.1

Ericaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted and thick-walled late-
wood fibers and by a lack of vessels
in terminal latewood. Wood semi-ring-
porous. Vessels predominantly solitary.
Mean tangential diameter of earlywood
vessels 20-50 μm, 100-200 vessels per
mm2. Fibers thin- to thick-walled. Axial
parenchyma diffuse in aggregates and
scanty paratracheal. Rays per mm 4-12.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits opposite and alternate, minute
(less than 4 μm in diameter). Vessels-
ray pits with distinct borders, similar
to intervessel pits in size and shape
throughout the ray cell. Earlywood ves-
sels length 200-500 μm. Fibers with
simple to minutely bordered pits (libri-
form fibers). Body ray cells procumbent
with 1-4 rows of square marginal cells.

›T ›T

Tangential section
Rays of two distinct sizes. Ray width
predominantly 1 to 3 cells. Larger rays
commonly 4 to 10 seriate.
Erica arborea L.

Wood basic density: 0,72 g/cm³

›T ›T

275
Erica sicula Guss. subsp. libanotica (Barb. Boiss. et
Barbey) P.F. Stevens
Ericaceae

PLANT DESCRIPTION
Evergreen shrub up to 65 cm high.
Leaves linear in whorls of 4. Native to
Cyprus, with limited distribution, grow-
ing in fissures of limestone rocks (250-
850 m alt.). It is also found in Sicily,
southern Turkey, Lebanon and Libya.
Erica sicula Guss. subsp. libanotica (Barb.Boiss. et Barbey) P.F. Stevens

TWIG ANATOMY
Bark
Some rays become dilated. Scleren-
chyma cells in phloem and in cortex.
Fibers in tangential rows and scattered
or irregularly dispersed. Cell content in
parenchyma cells. Epidermis distinct in
polarized light.

Xylem
As stem wood.

›T

Pith
Pith shape squared. Heterogeneous
pith. Thick-walled parenchyma cells
present. Cells dimorphic. Cell content
present (dark staining substances). Pits
in transverse and in longitudinal cell
walls. Vascular bundles clearly sepa-
rate to not distinct.

›T ›T

276
Stem xylem: 2 5 9 13 21 22 24 30 40.1 49 52.3 56 58 61 69 70 76 78 86 96 97 98 103 105 109 116.2
Twig bark: B6 B7 B12 B13 B28 B33 B34
Pith: P1.4 P3.3 P4 P5 P9 P9.1 P10.1 P10.2

Ericaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted and thick-walled late-
wood fibers and by a lack of vessels
in terminal latewood. Wood diffuse-
porous. Vessels predominantly solitary.
Mean tangential diameter of earlywood
vessels less than 20 μm, 40-100 ves-
sels per mm2. Tyloses with thin walls
common. Gums and other deposits in
heartwood vessels. Fibers thick- to very

Erica sicula Guss. subsp. libanotica (Barb.Boiss. et Barbey) P.F. Stevens


thick-walled. Axial parenchyma diffuse
to parenchyma scanty paratracheal.
Apotracheal parenchyma in narrow
bands or lines up to three cells wide.
›T ›T
More than 20 rays per mm.

Radial section
Simple perforation plates. Inter-vessel
pits opposite and alternate, minute
(less than 4 μm in diameter). Vessel-
ray pits with distinct borders, similar
to intervessel pits in size and shape
throughout the ray cell. Earlywood ves-
sel element length 100-200 μm. Fibers
with simple to minutely bordered pits
(libriform fibers). Uniseriate ray cells
square. Multiseriate rays with procum-
bent, square and upright cells mixed
throughout the ray.

›T ›T

Tangential section
Rays of two distinct sizes. Rays pre-
dominantly uniseriate. Larger rays
commonly 4 to 10 seriate.

Wood basic density: 0,75 g/cm³

›T ›T

277
Euphorbia hierosolymitana Boiss.
Woody Spurge
Euphorbiaceae

PLANT DESCRIPTION
Glabrous shrub up to 1 m high, with
milky latex. Leaves opposite, elliptic to
obovate. A very rare indigenous spe-
cies to Cyprus, occurring on dry slopes
in garigue and maquis (100-500 m
alt.). An eastern Mediterranean spe-
cies.

TWIG ANATOMY
Bark
Groups of sieve tube present. Sclereids
in tangential rows. Crystal sand pre-
sent. Cortex consist of large and small
parenchyma cells. Phellem homogene-
ous. Epidermis distinct in polarized
light.

Xylem
As stem wood.

›T
Euphorbia hierosolymitana Boiss.

Pith
Pith not visible in polarized light.
Pith shape round. Heterogeneous
pith. Unlignified cells present. Crys-
tal druses present. Pits in transverse
and in longitudinal cell walls. Vascular
bundles in the pith. Vascular bundles
clearly separate. Tracheary elements of
metaxylem in distinct radial rows.

›T ›T

278
Stem xylem: 2.1 5 7 10 13 20 21 24 32 40.1 50.1 52.3 61 69 70.2 75 96 105 116.2
Twig bark: B1 B4.1 B17 B23 B31 B33
Pith: P0.1 P1 P3.4 P6.2 P9 P9.1 P10 P10.1 P12

Euphorbiaceae
STEM ANATOMY
Transverse section
Only one ring in the observed sample.
Wood diffuse-porous. Vessels in radial
pattern. Vessels in radial multiples of 4
or more common. Mean tangential di-
ameter of earlywood vessels less than
20 μm, 100-200 vessels per mm2. Fib-
ers thin- to thick-walled. Tension wood
present. Axial parenchyma absent or
extremely rare or not to recognizable.
More than 20 rays per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits scalariform. Inter-vessel pits op-
posite, minute (less than 4 μm in
diameter). Vessel-ray pits with large
horizontal or vertical apertures. Early-
wood vessel element length 100-200
μm. Fibers with simple to minutely
bordered pits (libriform fibers). All ray
cells upright.

›T

Tangential section Euphorbia hierosolymitana Boiss.


Rays exclusively uniseriate.

Wood basic density: 0,33 g/cm³

›T ›T

279
Euphorbia thompsonii Holmboe
Thompson’s Spurge
Euphorbiaceae

PLANT DESCRIPTION
Robust, erect, tomentose perennial
herb with biennial flowering stems,
containing milky latex. A rare indig-
enous species occurring on dry slope
in garigue, maquis and pine forests
(50-400 m alt.). An east Mediterranean
species restricted to Cyprus and south-
ern Turkey.

TWIG ANATOMY
Bark
Groups of sieve tube present. Sclereids
scattered or irregularly dispersed.
Crystal sand present. With secretory el-
ements in ducts. Cell content in paren-
chyma cells. Phellem homogeneous,
distinct in polarized light.

Xylem
As stem wood.

›T
Euphorbia thompsonii Holmboe

Pith
Pith not visible in polarized light. Pith
shape round. Unlignified cells present.
Vascular bundles clearly separate to
not distinct.

›T

280
Stem xylem: 1 5 7 10 13 22 26 31 40.1 50.1 52.3 61 69 76 78 89 89.1 96 105 116.1
Twig bark: B1 B18 B23 B25 B28 B26 B31 B14.3
Pith: P1 P3.4 P10.1 P10.2

Euphorbiaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
zones with fibers of variable cell wall
thickness. Wood diffuse-porous. Ves-
sels in diagonal and/or radial pattern.
Vessels in radial multiples of 4 or
more common. Mean tangential diam-
eter of earlywood vessels less than 20
μm, 100-200 vessels per mm2. Fibers
thin- to thick-walled. Axial parenchy-
ma diffuse. Axial parenchyma scanty
paratracheal. Apotracheal parenchyma
in marginal or in seemingly marginal
bands, dark in polarized light. Rays 12-
›T ›T
20 per mm.

Radial section
Simple perforation plates. Inter-vessel
pits alternate, medium (7-10 μm in
diameter). Vessel-ray pits rounded or
angular with large apertures. Early-
wood vessel element length 100-200
μm. Fibers with simple to minutely
bordered pits (libriform fibers). All ray
cells upright and square.

›T ›T

Tangential section Euphorbia thompsonii Holmboe


Rays exclusively uniseriate.

Wood basic density: 0,38 g/cm³

›T ›T

281
Ricinus communis L.
Castor Oil Plant
Euphorbiaceae

PLANT DESCRIPTION
Deciduous shrub up to 5 m high, with
thick, hollow stems. A naturalised inva-
sive, widely distributed, mainly along
water canals and wastelands, especial-
ly in lowlands (0-500 m alt.). Probably
native to northeast tropical Africa, but
widely cultivated and naturalized in the
warmest part of the world.

TWIG ANATOMY
Bark
Collapsed sieve tubes present. Scleren-
chyma cells in phloem and in cortex.
Sclereids in tangential rows. Prismatic
crystals present. Cell content in paren-
chyma cells. Cortex consist of large
and small thin-walled parenchyma
cells. Phellem homogeneous, distinct
in polarized light.

Xylem
As stem wood.

›T

Pith
Pith shape round. Cells dimorphic.
Crystal druses present. Pits in trans-
verse and in longitudinal cell walls,
of two distinct size. Vascular bundles
clearly separate. Tracheary elements of
metaxylem in distinct radial rows.
Ricinus communis L.

›T ›T

282
Stem xylem: 2.1 5 9 9.1 13 21 25 30 41 49 53.1 60 61 67 69 70 96 97 105 116.1
Twig bark: B4 B7 B17 B20 B28 B31 B33
Pith: P1 P4.1 P6.2 P9 P9.1 P9.3 P10.1 P12

Euphorbiaceae
STEM ANATOMY
Transverse section
Only one ring in the observed sample.
Wood diffuse-porous. Vessels solitary
or in radial multiples of 2 to 4. Mean
tangential diameter of earlywood ves-
sel lumina 50-100 μm, 40-100 vessels
per mm2. Parenchyma-like fiber bands
alternating with ordinary fibers. Fibers
thick- to very thick-walled. Rays 12-20
per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits opposite, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 200-
500 μm. Vascular and/or vasicentric
tracheids present. Fibers with simple
to minutely bordered pits (libriform fib-
ers). All ray cells upright and square.

›T ›T

Tangential section
Rays uniseriate to 3 cells wide.
Ricinus communis L.

Wood basic density: 0,40 g/cm³

›T ›T

283
Alhagi graegorum Boiss.
Fabaceae

PLANT DESCRIPTION
A spinose subshrub, 30-60 cm high,
very similar to A. maurorum. Leaves
alternate, oblanceolate to ovate, thinly
hairy. An uncommon native to Cyprus,
growing mostly near coasts or a weed
in inland cereal fields (0-150 m). Na-
tive of east-Mediterranean countries
towards Iran.

TWIG ANATOMY
Bark
Sieve tube in clusters. Collapsed sieve
tubes present. Distinct rays dilatation.
Sclerenchyma cells in phloem and in
cortex. Fibers in small groups. Prismat-
ic crystals and crystal druses present.
With secretory elements in ducts. Cell
content in parenchyma cells.

Xylem
As stem wood.

›T

Pith
Pith shape round. Heterogeneous pith.
Thick-walled parenchyma cells. Cells
Alhagi graegorum Boiss.

dimorphic. Cell content present (dark


staining substances). Prismatic crystals
present. Pits in transverse and longi-
tudinal cell walls. Vascular bundles
clearly separate to not distinct. Axial
cells in regular rows (radial section).

›T ›T

284
Stem xylem: 1 5 9 9.1 13 22 25 30 41 45 49 52.3 56 58 61 67 69 78 96 97 102 103 105 116.2 136 136.1
Twig bark: B1 B4 B5 B7 B14 B20 B22 B25 B28
Pith: P1.2 P3.3 P4 P5 P6 P9 P9.1 P10.1 P10.2 P13

Fabaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
zones with fibers of variable cell wall
thickness. Wood diffuse-porous. Ves-
sels solitary and in radial multiples of
2 to 4. Mean tangential diameter of
earlywood vessel lumina 50-100 μm,
40-100 vessels per mm2. Vessels of
two distinct diameter classes. Tyloses
with thin walls common. Gums and
other deposits in heartwood vessels.
Parenchyma-like fiber bands alternat-
ing with ordinary fibers. Fibers thin- to
thick-walled. Axial parenchyma scanty
paratracheal. More than 20 rays per
›T ›T
mm.

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 100-
200 μm. Fibers with simple to minutely
bordered pits (libriform fibers). All ray
cells upright and square. Prismatic
crystals in ray parenchyma cells.

›T ›T

Tangential section
Rays of two distinct sizes: uniseriate an
2-3 cells wide. Larger rays height more
Alhagi graegorum Boiss.

than 1 mm.

Wood basic density: 0,55 g/cm³

›T ›T

285
Alhagi maurorum Medik.
Camel Thorn
Fabaceae

PLANT DESCRIPTION
A spinose subshrub, with glaucous,
glabrous shoots, pinkish flowers; fruit
a cylindrical, dry pod. An uncommon
native to Cyprus, occurring mostly
near coastal areas (0-150 m). Native of
east-Mediterranean countries towards
Iran and other Asian countries.

Photo: Hadjikyriakou
TWIG ANATOMY
Bark
Groups of sieve tube present. Distinct
rays dilatation. Sclerenchyma cells in
phloem and in cortex. Fibers grouped.
Prismatic crystals present. With large
laticifers, secretory elements, oil ducts
or mucilage ducts. Phellem homoge-
neous. Epidermis distinct in polarized
light.

Xylem
Vessels predominantly in clusters.
Rays usually not more than 3 seriate.

›T

Pith
Pith shape round to polygonal. Cells
dimorphic. Cell content present (dark
staining substances). Prismatic crystals
Alhagi mauroru Medik.

present. Pits in transverse and in lon-


gitudinal cell walls. Vascular bundles
clearly separate. Tracheary elements of
metaxylem in distinct radial rows.

›T ›T

286
Stem xylem: 2 5 9 9.1 13 22 25 30 41 45 49 52.3 61 67 69 78 79 83 96 97 103 105 116.1
Twig bark: B1 B6 B7 B14 B20 B25 B31 B33
Pith: P1 P1.1 P4.1 P5 P6 P9 P10.1 P12

Fabaceae
STEM ANATOMY
Transverse section
Growth ring indistinct or absent. Wood
diffuse-porous. Vessels solitary and in
radial multiples of 2 to 4. Mean tan-
gential diameter of earlywood vessel
lumina 50-100 μm, 40-100 vessels per
mm2. Vessels of two distinct diameter
classes. Fibers thin- to thick-walled.
Parenchyma-like fiber bands alternat-
ing with ordinary fibers. Axial paren-
chyma scanty paratracheal to vasicen-
tric, sometimes confluent. Rays 12-20
per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 100-
200 μm. Fibers with simple to minutely
bordered pits (libriform fibers). All ray
cells upright and square.

›T ›T

Tangential section
Rays of two distinct sizes: uniseriate an
2-3 cells wide.
Alhagi mauroru Medik.

Wood basic density: 0,38 g/cm³

›T ›T

287
Anagyris foetida L.
Bean Trefoil
Fabaceae

PLANT DESCRIPTION
Deciduous shrub up to 3 m high.
Leaves with 3 leaflets, flowers in ra-
cemes, yellow with brown blotches.
Native to Cyprus, usually occurring on
limestone slopes with shrubby vegeta-
tion (0-1200 m alt.). Indigenous also
in many other Mediterranean countries
and eastward to Iran and Arabia.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclerenchyma cells
in phloem and in cortex. Fibers and
sclereids in groups. Phellem homoge-
neous. Phellem and epidermis distinct
in polarized light.

Xylem
Mean tangential diameter of earlywood
vessels 20-50 μm. Earlywood vessels
length 200-500 μm. Rays predominant-
ly uniseriate, 12-20 rays per mm.

›T

Pith
Pith shape round. Thick-walled paren-
chyma cells present. Cells dimorphic.
Pits in transverse and in longitudinal
cell walls. Vascular bundles clearly sep-
arate. Tracheary elements of metax-
ylem in distinct radial rows. Axial cells
Anagyris foetida L.

›T
in regular rows (radial section).

›T

288
Stem xylem: 1 4 5 11 13 22 30 41 45 52.3 58 60 61 67 69 70.2 70.3 79 83 86 89 89.1 96 98 109 116.1
Twig bark: B1 B4 B7 B14 B19 B31 B33
Pith: P1 P3.3 P4 P9 P9.1 P10.1 P12 P13

Fabaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
radially flatted and thick-walled late-
wood fibers and by a lack of vessels in
terminal latewood. Wood semi-ring to
diffuse-porous. Mean tangential diam-
eter of earlywood vessel lumina 50-100
μm. Vessels of two distinct diameter
classes. Gums and other deposits in
heartwood vessels. Fibers thick- to
very thick-walled. Parenchyma-like
fibers alternating to ordinary fibers.
Paratracheal parenchyma confluent.
Apotracheal parenchyma in marginal or
in seemingly marginal bands, dark in
›T ›T
polarized light. Rays 12-20 per mm.

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in diame-
ter). Vessel-ray pits with distinct bor-
ders, similar to inter-vessel pits in size
and shape. Earlywood vessel element
length 100-200 μm. Vascular and/or
vasicentric tracheids present. Fibers
with simple to minutely bordered pits
(libriform fibers). Rays with procum-
bent, square and upright cells mixed
throughout the ray.

›T ›T

Tangential section
Rays commonly 4 to 10 seriate.
Anagyris foetida L.

Wood basic density: 0,58 g/cm³

›T ›T

289
Argyrolobium uniflorum (Dec.) Jaub. et Spach
Fabaceae

PLANT DESCRIPTION
Subshrub 5-20 cm high, with yellow
flowers; fruit a linear grey-brown pod.
A very rare native to Cyprus, recorded
in garigue vegetation in northern Cy-
prus (0-100 m alt.). Indigenous also to
north Africa, Syria, Palestine, Lebanon
and Turkey.

TWIG ANATOMY
Bark
Groups of collapsed sieve tube pre-
sent. Fibers scattered or irregularly
dispersed. Prismatic crystals present.
Epidermis distinct in polarized light.

Xylem
As stem wood.
Argyrolobium uniflorum (Dec.) Jaub. et Spach

›T

Pith
Pith shape triangular to roundish. Het-
erogeneous pith. Thick-walled paren-
chyma cells present. Cells dimorphic.
Pits in transverse and in longitudinal
cell walls. Vascular bundles clearly
separate to not distinct. Axial cells in
regular rows (radial section).

›T ›T

290
Stem xylem: 1 5 7 9.1 13 22 25 30 40.1 49 52.2 61 70 70.2 78 96 99 102 103 109 116.2 136 136.1
Twig bark: B4 B13 B20 B31 B33
Pith: P1.2 P3.3 P4 P9 P9.1 P10.1 P10.2 P13

Fabaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers and by a lack of vessels in ter-
minal latewood. Wood diffuse-porous.
Vessels in diagonal and radial pattern.
Vessels in radial multiples of 2 to 4
common. Mean tangential diameter
of earlywood vessels less than 20 μm,
40-100 vessels per mm2. Fibers very
thick-walled. Tension wood present.
Axial parenchyma scanty paratracheal.
More than 20 rays per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 50-
100 μm. Fibers with simple to minutely
bordered pits (libriform fibers). Rays

Argyrolobium uniflorum (Dec.) Jaub. et Spach


with procumbent, square and upright
cells mixed throughout the ray. Pris-
matic crystals in ray parenchyma cells.

›T ›T

Tangential section
Rays of two distinct sizes: uniseriate
and more than 10 seriate. Larger rays
height more than 1 mm.

Wood basic density: 0,67 g/cm³

›T ›T

291
Astragalus echinus subsp. echinus DC.
Fabaceae

PLANT DESCRIPTION
A much branched subshrub, up to 60
cm high. It occurs in two varieties:
chionistrae which is endemic to the
highest part of Troodos (1700-1900
m alt.), and echinus, which is locally
common at mid altitudes (400-1250 m
alt.). The latter is also indigenous to
Syria, Lebanon and Palestine.

TWIG ANATOMY
Bark
Groups of sieve tube present. Distinct
ray dilatation. Sclerenchyma cells in
phloem and in cortex. Sclereids in
tangential rows. Prismatic crystals
present. Phellem homogeneous and
layered.

Xylem
As stem wood.
Astragalus echinus subsp. echinus DC.

›T ›T

Pith
Pith not visible in polarized light. Pith
shape round. Cells dimorphic. Cell con-
tent, dark staining substances. Axial
cells in regular rows (radial section).

›T ›T

292
Stem xylem: 1 2 3 4 11 13 20 22 27 30 39.1 41 45 47 52.3 61 69 70 77 78 86 89 99 100.2 102 108 114 120
Twig bark: B1 B5 B7 B15 B17 B20 B31 B29
Pith: P0.1 P1 P4 P5 P13

Fabaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
marginal parenchyma bands. Wood
ring-porous to semi-ring-porous. Ves-
sel clusters common. Vessels cell
wall thick. Mean tangential diameter
of earlywood vessel lumina 50-100
μm. Vessels of two distinct diameter
classes, 5-20 vessels per mm2. Fibers
thick- to very thick-walled. Apotracheal
parenchyma diffuse in aggregates or in
narrow bands or lines up to three cells
wide. Paratracheal parenchyma scanty
paratracheal. Apotracheal parenchyma
in marginal or in seemingly marginal
›T ›T
bands. Less than 4 rays per mm.

Radial section
Simple perforation plates. Inter-vessel
pits scalariform to alternate, large
(more than 10 μm in diameter). Vessel-
ray pits with distinct borders, similar
to intervessel pits in size and shape
throughout the ray cell. Earlywood ves-
sel element length 100-200 μm. Fib-
ers with simple to minutely bordered
pits (libriform fibers). Body ray cells
procumbent with over 4 rows of square
marginal cells. Prismatic crystals in ray
parenchyma cells.

Astragalus echinus subsp. echinus DC.


›T ›T

Tangential section
Rays commonly more than 10 seriate.
Ray height more than 1 mm. Rays dis-
appear in polarized light. Axial paren-
chyma and vessel elements stories.

Wood basic density: 0,34 g/cm³

›T ›T

293
Calycotome villosa (Poir.) Link
Thorny Broom
Fabaceae

PLANT DESCRIPTION
Deciduous, spinose shrub up to 3 m
high. Leaves alternate, compound
with 3 narrowly obovoid leaflets; hairy
particularly below. Flowers yellow, fruit
an oblong, hairy pod. Native to Cy-
prus, locally very frequent (0-1000 m
alt.). Widespread in the Mediterranean
region.

TWIG ANATOMY
Bark
Groups of sieve tube present. Sclereids
scattered or irregularly dispersed. Pris-
matic crystals present. Cell content in
parenchyma cells. Phellem homogene-
ous.

Xylem
Vessels in short radial multiples. Mean
tangential diameter of earlywood ves-
sel lumina less than 20 μm.

›T
Calycotome villosa (Poir.) Link

Pith
Pith shape round. Cells dimorphic. Pits
in transverse and in longitudinal cell
walls. Vascular bundles clearly sepa-
rate. Axial cells in regular rows (radial
section).

›T ›T

294
Stem xylem: 1 1.1 4 8 10 11 13 22 26 30 36 39.1 40.2 45 48 52.3 58 60 61 70 77 79 83 97 103 109 115
Twig bark: B1 B15 B18 B20 B28 B31
Pith: P1 P4.1 P9 P9.1 P10.1 P13

Fabaceae
STEM ANATOMY
Transverse section
Growth rings boundaries clearly de-
marcated rings only along some ra-
dii. Wood semi-ring-porous. Vessels
in dendritic pattern. Vessels in radial
multiples of 4 or more common or in
clusters. Vessels cell wall thick (more
than 2 μm). Mean tangential diam-
eter of earlywood vessel lumina 20-50
μm. Vessels of two distinct diameter
classes, 20-40 vessels per mm2. Gums
and other deposits in heartwood ves-
sels. Fibers very thick-walled. Axial
parenchyma diffuse in aggregates and
vasicentric to confluent. Rays 4-12 per
›T ›T
mm.

Radial section
Simple perforation plates. Inter-vessel
pits alternate, medium (7-10 μm in
diameter). Vessels-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Helical thickenings in vessel elements
present. Earlywood vessel element
length 100-200 μm. Vascular and/or
vasicentric tracheids present. Fibers
with simple to minutely bordered pits
(libriform fibers). Rays with procum-
bent, square and upright cells mixed
throughout the ray.

›T ›T

Tangential section Calycotome villosa (Poir.) Link


Ray width predominantly 1 to 3 cells.
Rays of two distinct sizes.

Wood basic density: 0,61 g/cm³

›T ›T

295
Cercis siliquastrum L.
Judas Tree, Red Bud
Fabaceae

PLANT DESCRIPTION
Deciduous shrub or tree, up to 10 m
high, with broad crown, red-brown
branches and grey bark. Leaves alter-
nate, sub-orbicular to reniform, 6-13
cm in diameter. Native to southeast
Europe and western Asia, it has been
planted in many places (0-1500 m alt.).

Photo: Hadjikyriakou
TWIG ANATOMY
Bark
Collapsed sieve tubes present. Some
rays become dilated. Sclerenchyma
cells in phloem and in cortex. Fibers
scattered or irregularly dispersed, and
grouped. Fibers with an innermost
unlignified layer. Sclereids in groups.
Prismatic crystals. Phellem homogene-
ous.

Xylem
Mean tangential diameter of earlywood
vessel lumina 20-50 μm. Rays mostly
uniseriate.

›T

Pith
Pith shape round. Heterogenous pith.
Thick-walled parenchyma cells pre-
sent. Cells dimorphic. Cell content
present (dark staining substances). Pits
Cercis siliquastrum L.

in transverse and in longitudinal cell


walls. Vascular bundles clearly sepa-
rate. Tracheary elements of metaxylem
in distinct radial rows. Axial cells in
regular rows (radial section).

›T ›T

296
Stem xylem: 1 3 11 13 22 26 30 36 41 49 53.1 60 61 69 79 83 97 98 103 104 115 120 136 141.1
Twig bark: B4 B6 B7 B10 B9 B13 B14 B15 B19 B20 B31
Pith: P1 P3 P3.3 P4 P5 P9 P9.1 P10.1 P12 P13

Fabaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
the difference in vessel size between
latewood and earlywood and by ra-
dially flatted latewood fibers. Wood
ring-porous. Vessel predominantly in
clusters. Mean tangential diameter of
earlywood vessel lumina 50-100 μm,
40-100 vessels per mm2. Fibers thin- to
thick-walled. Axial parenchyma vasi-
centric, sometimes confluent. Rays per
mm 4-12.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, medium (7-10 μm in
diameter). Vessels-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Helical thickenings in vessel elements
present. Earlywood vessels length 200-
500 μm. Vascular and/or vasicentric
tracheids present. Fibers with simple
to minutely bordered pits (libriform fib-
ers). All ray cells procumbent. Prismat-
ic crystals in axial parenchyma cells.

›T ›T

Tangential section
Rays of to distinct sizes. Ray width
predominantly 1 to 3 cells. Larger rays
commonly 4 to 10 seriate. Axial paren-
chyma and vessel elements storied.
Cercis siliquastrum L.

Wood basic density: 0,61 g/cm³

›T ›T

297
Genista fasselata Decne
Thorny Gorse
Fabaceae

PLANT DESCRIPTION
Much branched shrub up to 3 m high,
with yellow flowers. Native to Cyprus,
occurring in two varieties: crudelis, en-
demic to Cyprus, confined to Troodos
(1200-1800 m alt.); fasselata with a
much wider distribution on the island
and it is also indigenous to the Aegean
islands, Lebanon and Palestine.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes in tangential rows. Some
rays become dilated. Sclerenchyma
cells in phloem and in cortex. Fibers
and sclereids in tangential rows. Ac-
icular crystals present. With secretory
elements in ducts. Phellem distinct in
polarized light.

Xylem
Wood diffuse-porous. Rays predomi-
nantly uniseriate. All ray cells upright
or square.
›T

Pith
Pith shape round. Cells dimorphic.
Crystal druses present. Pits in trans-
verse and in longitudinal cell walls.
Genista fasselata Decne

Vascular bundles clearly separate.


Axial cells in regular rows (radial sec-
tion). ›T

›T

298
Stem xylem: 1 4 7 11 13 22 25 30 36 39.1 40.2 45 49 52.3 58 60 61 67 69 70 70.3 78 97 103 106 115 119 120
Twig bark: B1 B2 B4 B6 B7 B12 B17 B21 B25 B26 B33
Pith: P1 P4 P6.2 P9 P9.1 P10.1 P13

Fabaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood semi-ring-porous. Vessels
in diagonal and/or radial pattern or in
clusters. Vessels cell wall thick (more
than 2 μm). Vessels of two distinct
diameter classes, mean tangential di-
ameter of earlywood vessels 20-50 μm,
40-100 vessels per mm2. Gums and
other deposits in heartwood vessels.
Parenchyma-like fibre bands alternat-
ing with ordinary fibers. Fibers thick-
to very thick-walled. Axial parenchyma
scanty paratracheal. Rays 4-12 per mm.
›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Helical thickenings in vessel elements
present. Earlywood vessel element
length 100-200 μm. Vascular and/or
vasicentric tracheids present. Fibers
with simple to minutely bordered pits
(libriform fibers). Parenchyma-like fibre
bands alternating with ordinary fibers.
Body ray cells procumbent with one
row of square marginal cells.

›T ›T

Tangential section
Ray width predominantly 1 to 3 cells.
Rays of two distinct sizes. Low rays
storied, high rays not storied. Axial pa-
Genista fasselata Decne

renchyma and vessel elements storied.

Wood basic density: 0,62 g/cm³

›T ›T

299
Glycyrrhiza glabra L.
Liquorice
Fabaceae

PLANT DESCRIPTION
Erect subshrub up to 1 m high, with
purplish flowers; fruit a brownish flat
pod. Leaves alternate, compound with
lanceolate leaflets. A naturalized spe-
cies, present in Cyprus as a relict of
cultivation (0-300 m alt.). Widespread
in the Mediterranean coastal countries.

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Some rays become
dilated. Sclerenchyma cells in phloem
and in cortex. Fibers irregularly dis-
persed or in groups. Prismatic crystals
present. With laticifers. Cell content in
parenchyma cells. Phellem homogene-
ous, distinct in polarized light.

Xylem
Vessels in radial multiples. Earlywood
vessel element length 100-200 μm.
More than 20 rays per mm.
›T

Pith
Pith shape round. Medullary sheath
present. Heterogenous pith. Few scat-
tered thick-walled parenchyma cells
present. Cell content present (dark
staining substances). Pits in transverse
Glycyrrhiza glabra L.

and in longitudinal cell walls. Vascular


bundles clearly separate from one oth-
er to not distinct. Tracheary elements
of metaxylem in distinct radial rows.

›T ›T

300
Stem xylem: 1 2 5 9 13 20 25 32 39.1 40.1 52.2 60 61 70 78 79 97 100.2 109 115 136
Twig bark: B1 B4 B7 B12 B18 B20 B34
Pith: P1 P3 P3.3 P3.4 P10.1 P10.2

Fabaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
zones with fibers of variable cell wall
thickness. Wood diffuse-porous. Ves-
sels predominantly solitary. Vessels
cell wall thick (more than 2 μm). Mean
tangential diameter of earlywood
vessels less than 20 μm. Fibers very
thick-walled. Axial parenchyma scanty
paratracheal to vasicentric. Rays 4-12
per mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits scalariform, small (4-7 μm in diam-
eter). Vessel-ray pits with large hori-
zontal or vertical apertures. Earlywood
vessel element length 50-100 μm.
Vascular and/or vasicentric tracheids
present. Rays with procumbent, square
and upright cells mixed throughout the
ray. Prismatic crystals present.

›T ›T

Tangential section
Rays commonly 1 to 3 seriate. Rays
disappear in polarized light.
Glycyrrhiza glabra L.

Wood basic density: 0,56 g/cm³

›T ›T

301
Hedysarum cyprium Boiss.
Cyprus’ Sainfoin
Fabaceae

PLANT DESCRIPTION
Erect or suberect subshrub up to 30
cm high. Leaves alternate, compound
with oblong leaflets. Flowers pinkish,
fruit a flat, one or two seeded pod. A
rare endemic species to Cyprus, oc-
curring on eroded limestone or sand-
stone slopes with garigue vegetation
(150-600 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Sclerenchyma
cells in phloem and in cortex. Fibers in
tangential rows. Sclereids scattered or
irregularly dispersed. Prismatic crystals
present.

Xylem
Mean tangential diameter of earlywood
vessel lumina 20-50 μm. Axial paren-
chyma scanty paratracheal. Parenchy-
ma-like fiber bands alternating with
ordinary fibers. Rays uni- or biseriate.
All cells upright or square.
›T

Pith
Pith not visible in polarized light.
Hedysarum cyprium Boiss.

Pith shape round. Heterogenous pith.


Thick-walled parenchyma cells present.
Unlignified cells present. Vascular bun-
dles clearly separate to not distinct.

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302
Stem xylem: 2 5 9 13 20 27 32 39.1 40.1 48 52.2 61 70 89 89.1 98 100.2 109 115 136
Twig bark: B1 B4 B7 B12 B18 B20
Pith: P1 P3 P3.3 P3.4 P10.1 P10.2

Fabaceae
STEM ANATOMY
Transverse section
Growth ring indistinct or absent. Wood
diffuse-porous. Vessels predominantly
solitary. Vessels cell wall thick (more
than 2 μm). Mean tangential diameter
of earlywood vessels less than 20 μm,
20-40 vessels per mm2. Fibers very
thick-walled. Axial parenchyma in
marginal or seemingly marginal bands,
dark in polarized light. Rays 4-12 per
mm.

›T ›T

Radial section
Simple perforation plates. Inter-vessel
pits scalariform, large (more than 10
μm in diameter). Vessel-ray pits with
large horizontal or vertical apertures.
Earlywood vessel element length 50-
100 μm. Fibers with simple to minutely
bordered pits (libriform fibers). Rays
with procumbent, square and upright
cells mixed throughout the ray. Pris-
matic crystals present.

›T ›T

Tangential section
Rays commonly 4 to 10 seriate. Rays
Hedysarum cyprium Boiss.
disappear in polarized light.

Wood basic density: 0,33 g/cm³

›T ›T

303
Hippocrepis emerus (L.) Lassen subsp. emeroides (Boiss. et Spruner) Las-
sen = Coronilla emerus L. subsp. emeroides (Boiss. et Spruner) Lassen
Fabaceae

PLANT DESCRIPTION
Deciduous, spinose shrub up to 3 m
high. Leaves alternate, compound, with
glabrous obovate leaflets. Flowers yel-
low, fruit an oblong, hairy pod. Native
to Cyprus, locally very frequent (0-
1000 m alt.). Widespread in the Medi-
terranean basin.
Hippocrepis emerus (L.) Lassen subsp. emeroides (Boiss. et Spruner) Lassen

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Distinct ray dilata-
tion. Sclereids scattered or irregularly
dispersed and in groups. Prismatic
crystals present. Phellem homogene-
ous. Epidermis distinct in polarized
light.

Xylem
Vessels in short radial multiples. Mean
tangential diameter of earlywood ves-
sel lumina less than 20 μm.

›T

Pith
Pith not visible in polarized light. Pith
shape round. Heterogeneous pith. Un-
lignified cells present. Cells dimorphic.
Pits in transverse and in longitudinal
cell walls. Pits in transverse cell walls.
Vascular bundles clearly separate.
Tracheary elements of metaxylem in
distinct radial rows.

›T ›T

304
Stem xylem: 1 5 7 13 22 25 30 36 40.2 45 49 52.3 58 61 67 70 76 79 89 96 97 109 116.1
Twig bark: B1 B4 B5 B18 B19 B20 B31 B33
Pith: P1.1 P1 P3.4 P4 P9 P9.1 P10.1 P12

Fabaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by ra-
dially flatted and thick-walled latewood
fibers. Wood diffuse-porous. Vessels
in diagonal pattern. Mean tangential
diameter of earlywood vessels 20-50
μm, 40-100 vessels per mm2. Vessels
of two distinct diameter classes. Gums
and other deposits in heartwood ves-
sels. Parenchyma-like fiber bands alter-
nation with ordinary fibers. Fibers very
thick-walled. Apotracheal parenchyma
diffuse, paratracheal parenchyma vasi-

Hippocrepis emerus (L.) Lassen subsp. emeroides (Boiss. et Spruner) Lassen


centric. Axial parenchyma in marginal
or in seemingly marginal bands. Rays
›T ›T
12-20 per mm.

Radial section
Simple perforation plates. Inter-vessel
pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Helical thickenings in vessel elements
present. Earlywood vessel element
length 100-200 μm. Fibers with simple
to minutely bordered pits (libriform
fibers). Rays with procumbent, square
and upright cells mixed throughout the
ray.

›T ›T

Tangential section
Rays uniseriate and 1 to 3 cells wide.

Wood basic density: 0,71 g/cm³

›T ›T

305
Ononis spinosa L. (Boiss.) subsp. leiosperma (Boiss.) Širj.
Spiny Resth-harrow
Fabaceae

PLANT DESCRIPTION
Erect, spinose subshrub up to 80 cm
high. Leaves alternate, compound, with
3 small, hairy, ovate leaflets with den-
tate margins. Indigenous to Cyprus,
locally common on moist, but also dry
and stony soils (0-1300 m alt.). It is
also found in other east Mediterranean
countries towards Iran.

TWIG ANATOMY
Bark
Ononis spinosa L. (Boiss.) subsp. leiosperma (Boiss.) Širj.

Groups of sieve tube and collapsed


sieve tubes present. Large rays be-
come dilated. Fibers in tangential rows.
Prismatic crystals present. Epidermis
distinct in polarized light.

Xylem
Rays 1-3 seriate. All ray cells upright
and square. More than 20 rays per
mm.

›T

Pith
Pith shape round. Cells dimorphic. Pits
in transverse and in longitudinal cell
walls. Vascular bundles clearly sepa-
rate. Tracheary elements of metaxylem
in distinct radial rows.

›T ›T

306
Stem xylem: 1 3 9 9.1 13 22 25 30 40.2 49 53.1 58 61 70 70.2 76 78 79 86 96 98 102 103 109 115 120 121 136 136.1 141.1
Twig bark: B1 B4 B6 B12 B20 B33
Pith: P1 P4.1 P9 P9.1 P10.1 P12

Fabaceae
STEM ANATOMY
Transverse section
Growth ring boundaries distinct by
the difference in vessel size between
latewood and earlywood, and by zones
with fibers of variable cell wall thick-
ness. Wood ring-porous. Vessels soli-
tary and in radial multiples of 2 to 4
common. Mean tangential diameter of
earlywood vessels 20-50 μm, 40-100
vessels per mm2. Gum and other de-
posits in hearwood vessels. Fibers very
thick-walled. Tension wood present.
Axial parenchyma diffuse and scanty
paratracheal to vasicentric. Apotracheal
parenchyma in narrow bands or lines
›T ›T
up to three cells wide. Rays 4-12 per
mm.
Radial section
Simple perforation plates. Inter-vessel

Ononis spinosa L. (Boiss.) subsp. leiosperma (Boiss.) Širj.


pits alternate, small (4-7 μm in di-
ameter). Vessel-ray pits with distinct
borders, similar to intervessel pits in
size and shape throughout the ray cell.
Earlywood vessel element length 200-
500 μm. Fibers with simple to minutely
bordered pits (libriform fibers). Rays
with procumbent, square and up-
right cells mixed throughout the ray.
Prismatic crystals in rays and in axial
parenchyma cells.

›T ›T

Tangential section
Rays of two distinct sizes: uniseriate
and 4 to 10 seriate. Larger rays height
more than 1 mm. Axial parenchyma
and vessel elements storied. Fibers
storied.

Wood basic density: 0,41 g/cm³

›T ›T

307
Robinia pseudoacacia L.
Black Locust
Fabaceae

PLANT DESCRIPTION
Deciduous tree up to 25 m high. Stip-
ules spinose. Leaves alternate, com-
pound with ovate leaflets. It is native
to the southeastern United States, but
has been widely planted and natural-
ized elsewhere in temperate hemi-
sphere (0-1800 m alt.).

TWIG ANATOMY
Bark
Groups of sieve tube and collapsed
sieve tubes present. Some rays become
dilated. Sclerenchyma cells in phloem
and in cortex. Fibers in tangential rows
and grouped. Sclereids in groups. Pris-
matic crystals present. Phellem homo-
geneous, distinct in polarized light.

Xylem
Vessels predominately in clusters.
Mean tangential diameter of earlywood
vessels 20-50 μm. Rays exclusively
uniseriate.
›T

Pith
Pith shape round. Prismatic crystals
present. Pits in transverse and in lon-
gitudinal cell walls. Vascular bundles
Robinia pseudoacacia L.

clearly separate. Tracheary elements of


metaxylem in distinct radial rows.

›T ›T

308
Stem xylem: 1 3 9 11 13 22 26 36 42 50.2 52.3 56 61 70 79 86 97 104 116.1 120 136
Twig bark: B1 B4 B6 B7 B9 B12 B14 B15 B19 B20 B31 B33
Pith: P1 P6 P9 P9.1 P10.1 P12

Fabaceae
STEM ANATOMY
Transverse section